The effects of intra- and inter-specific competition on the distribution of stocked juvenile Atlantic salmon, Salmo salar L., in relation to depth and gradient in an upland trout, Salmo trutta L., stream

The relative effects of inter- and intra-specific competition on the distribution of stocked salmon in relation to depth and gradient were investigated in an upland stream during two summer sampling periods. The stream was divided into two areas by an impassable fish barrier, and trout were removed from the upstream section prior to 2 years of salmon stocking. A small amount of trout re-immigration to the cleared area occurred. Under sympatric conditions in the downstream section the fry of both species were significantly more abundant in shallow water, whereas the yearling and older fish tended to inhabit the deeper, slow-flowing areas. Under near allopatric conditions in the cleared section salmon fry changed their distribution to include deeper areas, with shallow, fast-flowing water becoming the least preferred habitat. This trend for a wide distribution of salmon fry over all the available habitat in the upstream section was maintained in the second year when parr were present but trout remained at very low densities. It was therefore concluded that high inter-specific competition from trout was responsible for restricting the distribution of salmon fry to shallow habitat in the control area. Intra-specific competition from older salmon apparently only affected the growth and survival of salmon fry. The regulating mechanisms involved in these inter- and intra-specific effects are discussed in terms of competition for stream resources and predation. Recommendations based on the findings are made for stocking and habitat management of salmonid waters.     

Infestations by Gyrodactylus sp. of Atlantic salmon, Salmo salar L., in Norwegian rivers

Young Atlantic salmon, Salmo salar , brown trout, Salmo trutta , and Arctic char, Salvelinus alpinus , from eight rivers in North and Mid Norway were examined for Gyrodactylus. The fish were collected from 1975 to 1980. A Gyrodactylus salaris type was observed infecting salmon from six of these rivers. No trout or char were infected. A high frequency and intensity of infection of salmon were observed in all but one of the rivers surveyed. In the Saltdalselva, only one specimen of Gyrodactylus infecting one fish was observed. No subsequent mortality of salmon was observed in this river while there were signs of a high mortality of salmon in the other rivers. The salmon parr were more frequently attacked than the fry, the mortality of salmon seemed to have happened in most of the rivers one year after the first observations of Gyrodactylus were made. The mortality of salmon and the high infection rate of Gyrodactylus in these rivers appears unique, and as far as known to the authors there are no other described cases of mortality due to Gyodactylus in Atlantic salmon in natural waters. The reasons for the outbreak of Gyrodactylus in these rivers are not known. Two theories are discussed: one that the fish were weakened by environmental factors and the other that Gyrodactylus was introduced from some of the infected salmon hatcheries in Scandinavia.     

Infection of Atlantic salmon, Salmo salar L., by Gyrodactylus salaris, Malmberg 1957, in the River Lakselva, Misvær in northern Norway

Gyrodacrylus salaris was most probably introduced to the River Lakselva in 1975 through stocking of Atlantic salmon from an infected hatchery. The parasite population grew rapidly, and the parasite spread throughout the entire watercourse during the summer of 1976. This epidemic situation led to mortality among the young Atlantic salmon, and the density of salmon parr was heavily reduced from 1976 to 1977. The density of salmon parr has remained close to zero since then, while there are no apparent trends toward decrease or increase in the density of brown trout. In spite of the reduced density of young salmon, a new epidemic has developed each year among the few young 0+ and 1+ Atlantic salmon present in the river. Results from successive sampling during the summer of 1987, 1988 and 1989 indicate that most of the presmolt salmon are attacked during their first summer or autumn of life. The infection develops into an epidemic during the first autumn, winter or the next summer. The build-up of the parasite burden on the fish leads in turn to mortality. Norwegian Atlantic salmon probably have no resistance against G. salaris, since the parasite has recently been introduced to Norwegian rivers.     

Fluctuating recruitment and variable life history of migratory brown trout, Salmo trutta L., in a small, unstable stream

The recruitment dynamics and life history of migratory brown trout, Sulmo trutta L., were investigated in a small Baltic coast stream subject to recurring drought. Spawning males consisted of both mature male parr (101–206 mm t.l. ) and migrant males (205–780 mm t.l. ). Spawning females were all migrants which delayed maturity until reaching a significantly greater size on average (424–805 mm t.l. ) than migrant males. Male: female ratios were very high in spawning aggregations (9–12 males: 1 female) with males representing up to five year-classes or more. Gametes from several generations of males per spawning event may be important for maintaining the genetic viability of this population with few female spawners per year. The amount of spawning was dependent on precipitation just prior to and during the spawning period since migrants could not enter the stream under drought conditions. Migrants did not overwinter in the stream.
Drought also caused variable fry mortality following emergence in early summer. Recruitment of 0+ parr in autumn varied from c . 175 to 3000 during 3 years. Smolts were relatively young (ages 1–2) and small (≥8 cm), and were significantly longer on average than sibling parr. Yet emigration of 1-year-olds was not related to 0+ parr size the previous autumn because of overlapping growth rates.
Persistence of the migratory brown trout in this unstable environment may be the consequence of (i) life history adaptation (e.g. short freshwater residence of both juveniles and spawners), and (ii) a complementary set of individual life histories where variation in age of migrant spawners and the occurrence of mature male parr result in a stable spawner population despite inconsistent recruitment of migrants to the sea.     

Seasonal and spatial microhabitat selection and segregation in young Atlantic salmon, Salmo salar L., and brown trout, Salmo trutta L., in a Norwegian river

Seasonal microhabitat selection by sympatric young Atlantic salmon and brown trout was studied by diving. Both species, especially Atlantic salmon, showed seasonal variation with respect to surface and mean water velocities and depth. This variation is partly attributed to varying water flows and water temperatures. In winter the fish sought shelter in the substratum. A spatial variation in habitat use along the river due to different habitat availabilities was observed. Both species occupied habitats within the ranges of the microhabitat variables, rather than selecting narrow optima. It is hypothesized that the genetic basis allows a certain range to the behavioural response. Microhabitat segregation between the two species was pronounced, with brown trout inhabiting the more slow-flowing and partly more shallow stream areas. Atlantic salmon tolerated a wider range of water velocities and depths. Habitat suitability curves were produced from both species. It is suggested that habitat suitability curves that are based on observations of fish occupancy of habitat at median or base flow may not be suitable in habitat simulation models, where available habitat is projected at substantially greater water flows.     

Population changes after two years of salmon (Salmo salar L.) stocking in upland trout (Salmo tmtta L.) streams

The survival of salmon stocked in upland trout streams in the presence of salmon parr was found to be only about half the value recorded when trout alone made up the resident stock. Changes in the trout population were also recorded following the two years of salmon stocking, and these suggested that the presence of salmon parr may also influence trout fry survival. The findings are discussed in the context of habitat competition and total stream holding capacity.     

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.     

Redistribution of juvenile salmonid fishes after localized catastrophic depletion

Juvenile Atlantic salmon and brown trout were depleted at three sites ( c . 108–380 m²) of a natural stream during the summer months of 1991 and 1992. Local population changes and movements of fish marked in sections adjacent to each depleted area were monitored thereafter. There was very little movement of marked salmon parr into the central regions of the depleted areas following the immediate post-marking period. Upstream movement by young-of-the-year fish from high density sections in mid-late summer was noted for trout but not salmon. Unmarked 1-year-old salmon parr immigrated into depleted areas in June 1992, and the pattern of recolonization was consistent with migration upstream from the adjoining river. It is concluded that resident salmon were very strongly site-attached and resource tracking was of no functional significance as a compensatory mortality mechanism. The occurrence of a long distance migratory component in the population during early-mid summer indicates that this, rather than local resource tracking, constitutes a potential compensatory mechanism.     

Production of juvenile Atlantic salmon, Salmo salar L., and brown trout, Salmo trutta L., within different sections of a small enriched Norwegian river

Growth, density and production of juvenile Atlantic salmon and brown trout were studied in three different sections of the Kvassheimsåna River in south-western Norway from 1979 to 1983. Section 1. in the upper part of the river, is located above a waterfall impassable for migratory salmonids and is surrounded by grazing land. Sections 2 and 3, in the middle and lower parts of the river, are influenced by agricultural activity. Total nitrogen concentration varied between 250 and 1000 μg l ^?1 in section 1 and 1500 and 2500 μg l^?1 in sections 2 and 3. Total phosphorus (Tot-P) concentrations also increased with decreasing altitude: 19–46 μg l^?1 in section I and 31–101 μg l ^?1 in sections 2 and 3. The number of 0 + salmon in sections 2 and 3 varied between 30.1 and 167.8 specimens 100 m ^?2, with means 90.2 and 95.2 specimens 100 m ^?2:, respectively; the density of 1 + salmon, with mean values of 16.3 and 51.0 specimens 100m^?2 was significantly correlated with the original fry density. The growth rate of 0+ salmon was not inversely related to cohort density, but was significantly so for 1 + salmon. Mean annual salmon production in section 2 was 1595 g 100 m^?2 year ¹, and in section 3 was 841 g 100m^?2 year ¹. A logarithmic function gave the best curve fit between salmon production and mean annual biomass. Thus, production levelled off for the highest values recorded in section 2, and perhaps approached the carrying capacity of the stream. A multiple regression analysis showed that yearly variation in 1 + salmon density was the single factor accounting for most of the total variability in production (60%). Variation in water temperature and nutrient content were not significantly related to variation in fish production. Densities of brown trout were low in all sections (<20 specimens 100m ^?2). Fry density was highest in section 3 and parr density in section 1. All age groups of sympatric brown trout grew significantly faster in sections 2 and 3 compared with allopatric brown trout in section 1.     

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.     

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

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

High frequency of natural hybrids between Atlantic salmon, Salmo salar L., and brown trout, S. trutta L., in a Swedish river

Among 332 parr from the Swedish River Grönån examined by electrophoresis, 44 (13%) were hybrids between Atlantic salmon and brown trout. The hybrid frequencies in three sections of Grönån were significantly different (23. 8 and 2%). All hybrids are evidently of natural origin. and possible factors promoting hybridization are irregular overlapping spawning times. lack of separate spawning grounds, and involvement of sneakers.     

Natural hybrids between Atlantic salmon, Salmo salar L., and trout, Salmo trutta L., in juvenile salmonid populations in south-west England

Electrophoretic analysis of juvenile salmonids from south-west England revealed the presence of natural hybrids between Atlantic salmon, Salmo salar L., and brown trout, Salmo rrurra L., within three of five rivers. Amongst 409 juvenile 'salmon', 1.2% were hybrids, and of 150 juvenile 'trout' sampled, 2.0% were natural hybrids. In one river, similar hybridization levels in 'salmon' parr samples were found in two successive years. Levels of hybridization are high when compared with most similar studies, especially others within the United Kingdom. Possible factors influencing this are discussed.     

Temporal and spatial segregation of spawning in sympatric populations of Atlantic salmon, Salmo salar L., and brown trout, Salmo trutta L.

Based on data from Norwegian streams with sympatric populations of Atlantic salmon and brown trout, it is suggested that temporal segregation is the main mechanism segregating Atlantic salmon and brown trout during spawning. Peak spawning of trout was about 15 days earlier than that of salmon. Physical factors, such as water depth, water velocity and distance from the river banks segregate spawning sites of salmon and trout poorly. Gravel sizes of the redds of salmon and trout were significantly different, though with a considerable overlap, and mean egg depth of salmon and trout were 0.18 and 0.12 m, respectively, probably attributable to the different size of spawners of salmon and trout. None of the temporal or spatial parameters analysed segregate spawners of salmon and trout completely. Species determination of eggs and alevins from the redds showed no interspecific superimposition of redds. It is, therefore, concluded that low survival of hybrids after hatching does not explain the low frequency of hybrids observed in sympatric populations of salmon and trout.     

High mortality in introduced brown trout depressed potential gene flow to a wild population

Hatchery reared 0+ year brown trout Salmo trutta , with 51 mm mean L _T, were released in a sea trout stream in June 1991 to compare the survival of wild and introduced trout during the freshwater stage from age 0+ to 2+ years. The introduced brown trout were homozygous for a genetic marker, enabling released individuals and their offspring to be distinguished phenotypically from the local sea trout. The mean size of 0+ and 1+ year introduced parr was larger than 0+ and 1+ year wild parr, while 2+ year parr of both groups were of the same size. Half year survival rates of both introduced and wild parr increased with size up to c . 80 mm (1+ years), but then decreased, and could be described by a polynomial regression function, with the same shape for both groups. The introduced parr had, however, a significant lower survival rate than the wild parr. The number of the introduced cohort decreased from 2200 at release in 1991 to c . 20 in March 1994 ( c . 1% of the original number). Between 1994 and 2000 no introduced individuals or offspring have been observed in the study area. High mortality at the parr stage and additional mortality until the spawning, give a low probability for a genetic impact on the local population as long as releases are restricted, both in time and number of fish.     

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.     

The effects of intra- and inter-specific competition on the survival and growth of stocked juvenile Atlantic salmon, Salmo solar L., and resident trout, Salmo trutta L., in an upland stream

The relative effects of inter- and intra-specific competition on the survival and growth of stocked salmon were investigated in an upland trout stream during summer and winter sampling periods. The stream was divided into two areas by an impassable fish barrier, and trout were removed from the upstream section prior to 2 years of salmon stocking. Salmon fry stocked into the cleared area survived more than twice as well and grew significantly larger than those stocked into the area containing trout and older salmon. Intra-specific competition from older salmon in the second year of stocking in the cleared area significantly reduced the survival and growth of the O+ salmon. However, these were still significantly larger and survived better than those in the control area where inter-specific competition from trout was maintained. Some immigration of trout to the cleared area occurred; these showed greatly enhanced growth rates compared to those in the control area, reflecting low intra-specific trout competition in the former. Inter-specific competition effects of older salmon on both trout fry growth and survival were also detected, although the latter did not become apparent until the winter. This is discussed in terms of the relative importance of biotic and abiotic regulating mechanisms. Evidence of allopatric niche segregation is also discussed, since salmon in the cleared area did not have a biomass equivalent to that in the area which also contained trout.     

Lack of species-specific primer effects of odours from female Atlantic salmon, Salmo salar, and brown trout, Salmo trutta

We exposed, in two successive spawning seasons, individually placed precocious male Atlantic salmon ( Salmo salar ) and brown trout ( Salmo trutta ) parr to odour stimuli (ovarian fluid and urine mix) from ovulated conspecific or heterospecific anadromous females. Atlantic salmon parr had significantly higher plasma concentrations of the hormones 17α,20β-dihydroxy-4-pregnen-3-one (17,20β-P), 11-ketotestosterone (11-KT) and testosterone (T) after exposure to odours from conspecific females or from brown trout females compared to parr exposed to a control solution (0.9% NaCl). We did not observe any significant differences between the hormone levels in salmon parr exposed to the two female odours. The salmon parr exposed to conspecific odours had significantly higher volumes of strippable milt compared to the controls, but we did not find any significant differences when comparing the effect of the two female odours. Brown trout parr had significantly higher plasma 17,20β-P levels following exposure to heterospecific female odours compared to control males, but there was no significant difference between males exposed to the different female odours. We did not observe any significant differences in plasma levels of T and 11-KT and in milt volumes between exposed and control trout. Taken together, the results from both tested species indicate that the potency of heterospecific stimuli in stimulating increased plasma sex steroid hormone levels in male parr was as strong as stimuli from conspecific females. The results are discussed in connection to observed hybridisation between the two sympatric species.     

Otolith shape discriminates between juvenile Atlantic salmon, Salmo salar L., and brown trout, Salmo trutta L.

Otoliths of Atlantic salmon, Salmo salar L., are more slender than the otoliths of brown trout, Salmo trutta L. Discriminant analysis on otolith measurements of juvenile Atlantic salmon and brown trout from four river systems revealed a discriminant function which distinguished more than 94% of the cases. This function was tested by using data from a fifth river with cohabiting Atlantic salmon and brown trout: all Atlantic salmon and 91 % of the brown trout were correctly classified.     

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.