Above parr: Lowland river habitat characteristics associated with higher juvenile Atlantic salmon (Salmo salar) and brown trout (S. trutta) densities

Understanding juvenile salmonid habitat requirements is critical for their effective management, but little is known about these requirements in lowland rivers, which include important but unique salmonid habitats. We compared the relative influence of in-stream Ranunculus cover, water depth, prey abundance, distance upstream and two previously unexplored factors (water velocity heterogeneity and site colonisation potential) on summer densities of juvenile Atlantic salmon and brown trout. We applied electrofishing, habitat surveys and macroinvertebrate kick sampling, and calculated the site colonisation potential from salmon redd surveys across 18–22 sites in a lowland river in 2015–2017. Due to a recruitment crash in 2016, models including and excluding this unusual year were explored. Excluding 2016 data, juvenile salmon densities showed a positive association with Ranunculus cover and numbers of nearby upstream redds, and a negative association with distance upstream from the tidal limit. Trout densities were positively associated with velocity heterogeneity, indicating a potential indirect influence of Ranunculus mediated by water velocity. When including 2016, year had the largest effect on densities of both species, highlighting the impact of the recruitment failure. These findings uncover interspecific differences in the habitat requirements of juvenile salmonids in lowland rivers. Velocity heterogeneity and site colonisation potential had high explanatory power, highlighting that they should be considered in future studies of habitat use. These findings demonstrate that temporal replication and recruitment dynamics are important considerations when exploring species–habitat associations. We discuss potential management implications and argue that Ranunculus cover could be an important management tool in conservation of lowland salmonids.     

Spatial niche variability for young Atlantic salmon (Salmo salar) and brown trout (S. trutta) in heterogeneous streams

Abstract– Habitat is important in determining stream carrying capacity and population density in young Atlantic salmon and brown trout. We review stream habitat selection studies and relate results to variable and interacting abiotic and biotic factors. The importance of spatial and temporal scales are often overlooked. Different physical variables may influence fish position choice at different spatial scales. Temporally variable water flows and temperatures are pervasive environmental factors in streams that affect behavior and habitat selection. The more frequently measured abiotic variables are water depth, water velocity (or stream gradient), substrate particle size, and cover. Summer daytime, feeding habitats of Atlantic salmon are size structured. Larger parr (>7 cm) have a wider spatial niche than small parr. Selected snout water velocities are consistently low (3–25 cm. s^?1). Mean (or surface) water velocities are in the preferred range of 30–50 cm. s^?1, and usually in combination with coarse substratum (16–256 mm). However, salmon parr demonstrate flexibility with respect to preferred water velocity, depending on fish size, intra- and interspecific competition, and predation risk. Water depth is less important, except in small streams. In large rivers and lakes a variety of water depths are used by salmon parr. Summer daytime, feeding habitat of brown trout is also characterized by a narrow selection of low snout water velocities. Habitat use is size-structured, which appears to be mainly a result of intraspecific competition. The small trout parr (<7 cm) are abundant in the shallow swift stream areas (<20–30 cm depths, 10–50 cm. s^?1 water velocities) with cobble substrates. The larger trout have increasingly strong preferences for deep-slow stream areas, in particular pools. Water depth is considered the most important habitat variable for brown trout. Spatial niche overlap is considerable where the two species are sympatric, although young Atlantic salmon tend to be distributed more in the faster flowing and shallow habitats compared with trout. Habitat use by salmon is restricted through interspecific competition with the more aggressive brown trout (interactive segregation). However, subtle innate differences in behavior at an early stage also indicate selective segregation. Seasonal changes in habitat use related to water temperatures occur in both species. In winter, they have a stronger preference for cover and shelter, and may seek shelter in the streambed and/or deeper water. At low temperatures (higher latitudes), there are also marked shifts in habitat use during day and night as the fish become nocturnal. Passive sheltering in the substrate or aggregating in deep-slow stream areas is the typical daytime behavior. While active at night, the fish move to more exposed holding positions primarily on but also above the substrate. Diurnal changes in habitat use take place also in summer; brown trout may utilize a wider spatial niche at night with more fish occupying the shallow-slow stream areas. Brown trout and young Atlantic salmon also exhibit a flexible response to variability in streamflows, wherein habitat selection may change considerably. Important topics in need of further research include: influence of spatial measurement scale, effects of temporal and spatial variability in habitat conditions on habitat selection, effects of interactive competition and trophic interactions (predation risk) on habitat selection, influence of extreme natural events on habitat selection use or suitability (floods, ice formation and jams, droughts), and individual variation in habitat use or behavior.     

Selective segregation in intraspecific competition between juvenile Atlantic salmon (Salmo salar) and brown trout (Salmo trutta)

Interactive segregation has been suggested as the ruling competition mechanism determining niche and niche segregation between juvenile Atlantic salmon (Salmo salar) and brown trout (Salmo trutta). Results from allopatry–sympatry observations of habitat use in both nature and in experiments were contrary to predictions derived from the interactive segregation hypothesis. Habitat use parameters under natural conditions such as distance to shore for Atlantic salmon parr were nearly identical in allopatric (mean ± SD; 3.2 ± 1.4 m) and sympatric (3.3 ± 1.4 m) situations. Occupied water depths largely reflected available water, but water depths <15 cm were avoided by salmon parr. Under experimental conditions, habitat use of allopatric salmon was density independent and salmon size had only minor effects, with smaller fish being more likely to occur in the shallow. Habitat use of salmon in sympatry with trout did not differ from allopatric salmon habitat use, and only salmon size had minor effects on depth choice – occurrence of trout or fish density had no effect. Allopatric trout was in general more frequent in the shallow habitat than salmon. Habitat use of sympatric trout was affected by the occurrence of salmon and trout size, resulting in a higher use of the shallow habitats for small trout. To conclude, selective segregation has a dominant role in salmon habitat use (not affected by trout occurrence), whereas a mixed situation occurs in trout habitat use with elements of interactive segregation when competing with Atlantic salmon (affected by salmon occurrence).     

Morphological variability of Atlantic salmon Salmo salar and brown trout Salmo trutta in different river environments

Abstract – The morphologies of sympatric juvenile Atlantic salmon Salmo salar parr and brown trout Salmo trutta parr were compared between Irish rivers with contrasting hydraulic environments – a high‐gradient and a low‐gradient tributary from the River Barrow catchment, south‐east Ireland and a high‐gradient river from the Burrishoole catchment, west of Ireland. The two catchments differ markedly in mean annual precipitation (849.0 mm year^?1 and 1370.3 mm year^?1 for the Barrow and Burrishoole catchments, respectively). Parr of both species demonstrated morphological variation between and within catchments. Changes in metrics such as pectoral fin length, body depth and body length between and within catchments suggest that hydraulic forces were a major determinant of morphological variation. Both species from the Burrishoole catchment had relatively larger pectoral fins, longer heads, larger eyes and shallower bodies than conspecifics from the Barrow catchment. In rivers subject to frequent rainfall‐driven high discharges, such as those in the Burrishoole catchment, a more fusiform body and head shape coupled with larger pectoral fins may reduce the energetic cost of maintaining position in the water column, as well as increase stability and manoeuvrability. The larger eyes in both parr species in the Burrishoole catchment could further be a response to the lower visibility of the more turbid and peat‐stained waters or to the reduced prey availability. The results of this study demonstrate that local adaptation to flow conditions can yield morphologically distinct populations and that multiple species can exhibit parallel phenotypic responses to changing environmental conditions.     

Saltwater performance of triploid Atlantic salmon Salmo salar L. brown trout Salmo trutta L. hybrids

Survival and growth in a saltwater net-pen of sexually immature triploid Atlantic salmon Salmo salar L. brown trout Salmo trutta L. hybrids was comparable to that of immature diploid Atlantic salmon. Following 17 months of freshwater rearing, the experimental fish were individually tagged and transferred to a saltwater net-pen where they were raised communally for 376 days. Initial and final average weights were 158 and 760 g per fish for the diploid Atlantic salmon and 209 and 1010 g per fish for the triploid hybrids; weights for the hybrids were significantly larger in both cases (α= 0.05). Survival from transfer to harvest was 43% for the Atlantic salmon and 48% for the hybrids. Average specific growth rate of fish which survived to harvest was 0.42% day^-1 for Atlantic salmon and 0.41% day^-1 for hybrids; these values were not significantly different. No significant differences were observed in average condition factor and dress-out percentage between crosses. Average gonadal weights and gonadosomatic indices were not significantly different for male diploid Atlantic salmon, female triploid hybrids and male triploid hybrids, but were significantly greater for female Atlantic salmon.     

Magnesium requirement of Atlantic salmon (Salmo salar L.) parr in seawater-treated fresh water1

This experiment was undertaken to establish the magnesium (Mg) requirement in young Atlantic salmon, Salmo salar L., in seawater-treated fresh water. In Norwegian hatcheries it is a common practice to add sodium hydroxide and/or sea water (1–2%) to improve pH and conductivity of the natural fresh water. Parr with initial weight of 8 g, were divided in six triplicate groups in brackish water containing 54 mg Mg L^?1 and fed a basal casein-gelatine diet supplemented with minor amounts of krill and fish meal (containing 200 mg Mg kg^?1) for an initial period of 3 weeks. Thereafter the fish were fed this diet supplemented with either 0, 100, 200, 300, 400 or 500 mg Mg kg^?1 (as MgSO₄) for 12 weeks. Growth and feed efficiency were recorded. Concentrations of Mg and other divalent cations (Ca and Zn) were measured in whole fish, serum and vertebrae. Sodium concentration in vertebrae was also measured. Growth and feed efficiency were unaffected by the levels of dietary magnesium used in the experiment. Magnesium concentrations in the whole body, serum and vertebrae Mg appeared to be more sensitive than growth and feed efficiency to differences in dietary Mg intake. The group fed the unsupplemented diet showed significantly lower Mg concentration in these tissues than the other groups. Whole-body calcium concentration was negatively correlated with dietary Mg and Ca:Mg ratios in the vertebrae were significantly affected by the dietary Mg levels. Zinc concentration in whole body, serum and vertebrae was not altered by the dietary Mg levels. Further, vertebral Na concentration did not vary between the dietary treatments. In conclusion, a minimum Mg supplementation level of 100 mg kg^?1 dry diet (in total, 326 mg kg^?1) was needed to maintain Mg concentration in the whole body and serum and for proper bone mineralization.     

Swimming performance of wild and F1-hatchery-reared Atlantic salmon (Salmo salar) and brown trout (Salmo trutta) smolts

Abstract –  The swimming performance of wild and hatchery-reared smolts of two salmonid species was investigated. Wild Atlantic salmon smolts (WS) and brown trout smolts (WT) of equal size were caught in fish traps during migration. Hatchery-reared smolts of both species (HS and HT for salmon and trout respectively) were first generation offspring from wild broodstock. The swimming performance of individual smolts from the four groups (WS, HS, WT, HT) was tested three consecutive times using a swimming flume with water flowing at a start rate of 0.16 m·s⁻¹ and a constant acceleration rate of 0.167 cm·s⁻² (10 cm·s⁻¹·min⁻¹). Wild caught smolts of both species performed significantly better than those reared in hatchery conditions. The WS group were observed to maintain an average swimming speed ( U _burst) that was 30% faster than the HS group, whereas the wild trout smolts were superior to HT by approximately 25%. Repeated measures revealed species-specific exhaustion patterns. Brown trout smolts maintained consecutive U _burst indicating significant stamina compared with Atlantic salmon smolts that were found to be exhausted by the initial trial.     

River temperature and adult anadromous Atlantic salmon,Salmo salar,and brown trout,Salmo trutta

Abstract In terms of the spawning migration of adult salmon, Salmo salar L., water flow is often considered the primary factor controlling river entry and fluctuations in flow controlling when the fish subsequently migrate upstream. However, water temperature has also been suggested to modify the spawning migration of salmon, particularly their movements within estuaries and the timing of freshwater entry. Freshwater temperature is more likely to impact salmonid biology than flow, particularly in relation to temperature dependant metabolic costs, time of spawning and fecundity. Therefore, temperature may be more of a factor regulating salmonid populations in fresh water than flow itself. This study focuses on two aspects of the impact of temperature on salmonids in fresh water: first, how salmon may modify their behaviour to adapt to changes in temperature and second the potential relationship between temperature, environmental conditions (e.g. water quality) and physiology (e.g. maturation and olfaction) in regulating adult migration.     

Timing of spawning in cultured and wild Atlantic salmon (Salmo salar) and brown trout (Salmo trutta) in the River Vosso, Norway

Spawning time and size of wild and nonnative cultured female Atlantic salmon (Salmo salar L.) were studied in the River Vosso, Norway. Cultured immigrants were smaller, and peak spawning occurred 21 and 26 days earlier relative to wild salmon in 1991 and 1992, respectively. It was calculated that hatching and initial feeding of offspring of cultured females peaked 12 and 9 days earlier in spring 1992 and 19 and 8 days earlier in spring 1993 relative to offspring of wild females. Spawning time of the cultured salmon is similar to most Norwegian salmon populations and to cultured fish in other streams. In the River Vosso it also overlaps with that of brown trout (Salmo trutta L.), which may increase hybridization. The early time of breeding in nonnative fish indicates local adaptation in this trait in wild salmon. The consequences of early breeding of cultured fish are unpredictable in relation to spring temperatures but probably reduce their relative breeding success. The smaller size may also reduce the success of cultured females.     

The effects of ration size on migration by hatchery‐raised Atlantic salmon (Salmo salar) and brown trout (Salmo trutta)

Abstract – The possibility to increase the proportion of migrating hatchery‐reared smolts by reducing their food ration was studied. Lake‐migrating, hatchery‐reared salmon (Salmo salar) and trout (Salmo trutta) smolts were either fed normal rations, based on recommendations from the fish‐farming industry, or reduced (15–20%) rations. They were released into the River Klarälven, western Sweden, and followed as they swam downstream to Lake Vänern, a distance of around 25 km. For both Atlantic salmon and brown trout, smolts fed a reduced ration migrated faster than fish fed a normal ration. Furthermore, a higher proportion of salmon smolts fed reduced rations migrated to the lake than fish fed normal rations in 2007 but not in 2006. This difference between years corresponded to greater treatment differences in size and smolt status in 2007 than in 2006. For trout, the proportion of migrating individuals and smolt development did not differ with ration size. Trout migrants fed a normal ration had a higher standard metabolic rate (SMR) than nonmigrants, whereas there was no difference in SMR between migrating and nonmigrating salmon. These results show that it is possible to use a reduced food ration to increase the migration speed of both Atlantic salmon and brown trout and to increase the proportion of migrating Atlantic salmon.     

Population decline in the endemic Atlantic salmon (Salmo salar) in Kapisillit River,Greenland

Atlantic salmon, Salmo salar L., is found throughout the North Atlantic, with thousands of rivers having spawning populations. In Greenland, spawning is limited to one river in West Greenland, the Kapisillit River (64?N), and the salmon are limited to the lower few kilometres of the system. Using mark–recapture, it was estimated the parr population was 5,953 individuals, and that the population size has declined by 52% since 1959. In spite of this decline, parr density remains high, being between 0.26 and 0.62 parr/m². Using a historical age‐length key, an estimated minimum of 635 smolt will have descended to the sea in 2017. These will be caught in a fishery currently subject to no regulatory measures and fishing remains the most likely driver of the population decline. The genetically distinct population is endemic to Greenland, and managers should implement measures to conserve this genetic integrity and local biodiversity.     

Density of brown trout, Salmo trutta L., and Atlantic salmon, Salmo salar L., parr after point and scatter stocking of fry

Atlantic salmon, Salmo salar L., and brown trout, Salmo trutta L., fry were point and scatter stocked in the early part of June at densities of 63–263 fry 100 m⁻² per species in the River Viantienjoki, a small river in northern Finland, and their population densities were assessed in late summer. Both species were always stocked together in similar quantities. Point stocking was used in the first 2 years and scatter stocking in the following 2 years. In point stocking, there was no correlation between the distance from the stocking sites (maximum = 250 m) and parr density in census sites ( r = −0.013 and 0.019 for brown trout and Atlantic salmon, respectively). The stocking density of fry did not influence parr density in August by either method or by species. Stocking density explained only from 11% to 23% of the parr survival depending on the species or stocking method. The mean densities of Atlantic salmon and brown trout parr did not differ significantly from each other at any fishing site ( P > 0.05). Both point and scatter stocking appear to be suitable methods for use in small rivers. The parr densities depend more on the other factors (e.g. habitat quality) than the stocking method, and the choice between methods could be based on the time and labour available.     

Production of Atlantic salmon (Salmo salar) in fresh water and at sea at hgh and low densities

A basic management objective is to identify the production capacity of Atlantic salmon in both freshwater and marine environments. Production in the biological sense is the total elaboration of fish biomass over time, and it is a function of fish abundance, growth and survival. In this article, we consider the factors that affect production at low and high stock levels. Using data from New Brunswick (Miramichi and Restigouche Rivers) and Newfoundland (Western Arm Brook), we consider freshwater and marine production of salmon at low and high stock levels. For all life stages, freshwater and marine production are related to initial biomass of the year-class: thus, it is important to maintain adequate egg deposition. Production at high stock levels is stock-dependent and mortality is extremely variable, particularly in the marine environment.     

Incidence of hybridization between Atlantic salmon, Salmo salar L., and brown trout, Salmo trutta L., in Ireland

The substantial growth of the farmed salmon industry in Europe since the 1970s has highlighted concerns regarding the genetic impact of escaped farmed salmon on wild salmonid stocks. High incidences of salmon × trout hybrids have been recorded in rivers situated near intensive salmon farming in Norway and Scotland, which may be indicative of a breakdown in reproductive isolation between salmon, Salmo salar L., and brown trout, Salmo trutta L. In the present study, salmonid fry and 0+ parr were collected from rivers in western Ireland. Allozyme and minisatellite DNA analysis were carried out on fry to determine the frequency of F1 hybrids from 10 rivers located within 38 km of salmon farms and three rivers at least 80 km from salmon farms. A total of 49 hybrids were recorded from 4135 salmonid fry (frequency = 1.2%). Mitochondrial DNA analysis showed that all hybrids arose from Atlantic salmon female × brown trout male crosses. Hybrid parr were recorded from one of the low-risk rivers (1.0%), but were present in seven out of the 10 catchments located within 38 km of salmon farms, with frequencies ranging from 0.7% to 3.1%. The results of the present survey, which represents the first extensive record of the levels of salmon-trout hybridization in Ireland, are discussed in relation to the continued growth of salmon farming in this country.     

Control of phosphorus homeostasis of Atlantic salmon (Salmo salar) in fresh water

Studies were conducted to determine the absorption, excretion and requirement of dietary phosphorus (P) by Atlantic salmon (Salmo salar). Triplicate groups of salmon parr, initial weight 15 ± 0.5±g , were fed, diets containing 4, 5, 6, 7, 8, 9, 11 and 13 mg P and 20 KJ of digestible energy (DE) per±g of diet (dry matter basis, DMB) to satiation for 16 weeks. The basal diet containing 4 mg P g^-1 (0.15 mg available P per KJ DE) was supplemented with graded levels of calcium phosphate, Ca(H₂PO₄)²H₂O to formulate the eight experimental diets. The fish were reared in fresh water at a temperature of 15 °C on a 12 h photoperiod. Vertebrae ash increased from 316 to 516 mg g^-1 fat-free dry matter with an increase in dietary P content. P requirement was estimated by using a four-parameter sigmoidal equation. The data suggests that a diet of 0.28 mg available P per KJ DE is needed for Atlantic salmon. Phosphate and calcium levels in plasma and bone increased, whereas levels of magnesium and liver cholecalciferol decreased, with an increase in dietary P.Phosphate excretion in urine and apparent availability of P were determined in deficient and replete fish. In deficient fish, the urine phosphate concentration was 0.10 mmol L^-1 before feeding and 0.25 mmol l^-1 after feeding, whereas in replete fish these concentrations were 1.09 and 5.11 mmol l^-1, respectively. The increase in urine phosphate concentration was higher in replete fish than in deficient fish, however, the apparent absorption of P was found to be significantly lower in replete fish than in deficient fish. These results suggest that similarly to terrestrial vertebrates, P homeostasis in Atlantic salmon is controlled by absorption in the intestine, conservation in the kidney and storage in the bones.     

Effect of sudden increase in discharge in a large river on newly emerged Atlantic salmon (Salmo salar) and brown trout (Salmo trutta) fry

Abstract– Downstrcam displacement of salmonid fry due to flow increase, from 12–15 m³/_s to >100 m³_s, was documented in the river Suldalslåen, Western Norway. In May only fry of brown trout ( Sulmo trutta ) occurred in the drift, while from the beginning of June only newly emerged Atlantic salmon ( Sulmo sular ) were found. The maximum number of Atlantic salmon fry drifting during a single day was estimated to be 17 000 individual. Their density in the drift was higher during the night than during the day, and their appearance in the drift coincided with the predicted period of emergence. Total brown trout numbers in the drift were estimated to vary between 4000 and 16 000 per day. Fry displaced downstream from the lowermost part of the river were lost from the population. The total losses were estimated to be between 75 000 and 100 000 Atlantic salmon fry which represents between 5.6 and 11.1%) of juvenile mortality during the first year of life.     

Side-aspect target strength of Atlantic salmon (Salmo salar), brown trout (Salmo trutta), whitefish (Coregonus lavaretus), and pike (Esox lucius)

The side-aspect acoustic target strengths (TS) of 19 Atlantic salmons (Salmo salar), 16 brown trouts (Salmo trutta), 10 whitefish (Coregonus lavaretus) and 9 pikes (Esox lucius) were measured using a 200 kHz split-beam echosounder, in order to study the relationship between TS and fish size indices (length, weight and side area). The effect of side aspect angle on TS was also studied. Linear models between TS and the logarithm of the fish size indices were fitted with length being best for predicting TS. Typically, the standard error of estimate was 1.2–2.9 dB. The side-aspect TS measurements with specimens of known size showed that the linear relationship between full side-aspect TS and the logarithm of fish length for salmonid (Salmo salar + Salmo trutta) was on average 4.7 dB (SE = 0.7), lower than for whitefish and pike combined. The effect of side aspect angle on TS was modelled with cos³ (2α) function. The differences in the TS between full side aspect and head/tail aspect were 17.4, 19.0 and 19.6 dB for salmonid, whitefish, and pike, respectively.     

Within-stock variations of life-history traits in juvenile Atlantic salmon (Salmo salar)

Variations in space (geographical location) and time (year) of phenotypic traits linked to life-history (size, growth, condition and sexual maturation) were analysed within the population of Atlantic salmon (Salmo salar L.) 1 + parr of Little Codroy River (southwest Newfoundland). The hydrographic system was divided into 10 zones: 4 for the mainstem and 6 for the tributaries. Despite the small size of the river studied, a highly significant spatial heterogeneity was observed. Within the mainstem, size, growth and rate of maturation of males tended to decrease when progressing upstream. When compared with the tributaries as a whole, the mainstem had smaller salmon juveniles, both at the end of the first winter (mainstem: 67.6 mm; tributaries: 73.3 mm) and at 1 + age (mainstem: 80.6 mm; tributaries: 88.3 mm), and the proportion of maturing fish among 1 + males was lower (mainstem: 48.6%; tributaries: 74.2%). According to the feature considered, from 20% to 70% of the spatial effect was due only to differences between the mainstem and the tributaries as a whole. A strong year effect was also revealed. Life-history traits seemed to fluctuate over time independently from one tributary to another, whereas patterns in their yearly variations were basically consistent among zones in the mainstem. The potential role of genetic and environmental factors in explaining changes in space and time of biological characteristics of parr is discussed. Within a small system such as Little Codroy River, spatio-temporal life-history variations of Atlantic salmon juveniles were most likely driven by environment.