Sea trout in North Argyll Sea lochs, population, distribution and movements

Sea trout were sampled by shore seining in the sea lochs of the west coast of Scotland between 1970 and 1974, This area is of special interest because of the indentation of the coastline and the varying degree of freshwater and marine influence in the sea lochs. The majority of fish caught were in their first year after smolt migration. These post-smolts were caught mainly in May and June, after which the number of trout present was low until fish at the 'whirling' stage appeared in the catches at the end of August, continuing through to the following spring. An additional recruitment of mainly unsilvered young trout from the rivers to the sea lochs was found in the autumn. Age and sex composition of both spring and autumn recruits, and of mature fish, were investigated and compared.
A total of 3228 sea trout were tagged, with 311 recaptures. These recaptures supported the evidence from smolt trapping and beach seine catches that, in the first post-migration year, the smolts migrate from the rivers from late March to early May, they then migrate from the sea lochs in May and June and return in late summer and autumn.     

The growth of juvenile Atlantic salmon, Salmo salar L., and brown trout, Salmo trutta L., in a Scottish river system subject to cooling-water discharge

The River Fiddich, a tributary of the R. Spey in north-east Scotland, is a spawning river for both Atlantic salmon and brown trout. Warm cooling water effluent is discharged from several distilleries at different points in the lower reaches and raises the temperature of the river 1–3°C above ambient for most of the year. Salmon and trout grow more rapidly in this region than further upstream, and juvenile salmon generally migrate a year earlier, as 2 + smolts. Available data were too few to determine whether there was a similar difference for trout. Similar studies on the R. Dullan, a tributary of the Fiddich, and on the Cromdale Burn in the same area, confirmed that the growth rate of fish is faster downstream from distillery discharge points. It is suggested that increased invertebrate production may influence the growth rate.     

Temporal and spatial relationships in the movements of loch dwelling brown trout, Salmo trutta L., recorded by ultrasonic tracking for 24 hours

Ultrasonic telemetry was used to record simultaneously, over one complete April day, the movements of four adult brown trout, Salmo trutta , which were residents of a shallow Scottish loch. The daily movements were generally along linear tracks. The home ranges are described as standard and 95% confidence ellipses. The area of standard ellipses ranged from 330 to 633 m² while 95% confidence ellipses were larger, ranging from 513 to 979 m². The distances separating individuals, which ranged from 1 to 70 m, were greatest during peaks of activity. Although there was considerable spatial overlap of individual home ranges, the trout were not within visual range for most of the tracking period. Close encounters (proximity <2.5m) occurred most frequently and lasted longest at night. It is concluded, from this study, that brown trout in small productive lochs are solitary but may hold temporary station within widely overlapping home ranges.     

Conservation and management of brown trout, Salmo trutta, in Scotland: an historical review and the future

SUMMARY. 1. Intensive research into the life history of brown trout started In 1948 when the Brown Trout Laboratory was opened in Pitlochry. Over the next 15 years significant contributions were made to the brown trout literature upon which the Laboratory based advice to landowners and anglers wanting to develop their fisheries. 2. Increasing pressure from the government for more work on Atlantic salmon tended to divert research funds and time away from brown trout investigations. The International Biological Programme's major study at Loch Leven from 1966 to 1972 ensured a continuing interest in trout in standing waters. Over this period little attention had been paid to trout in rivers. This changed when a number of investigations were started on the River Tweed by Edinburgh University. 3. A major constraint to brown trout conservation and management has been illegal fishing and lack of records on stocking activities and catches. The granting of Protection Orders under the Freshwater and Salmon Fisheries (Scotland) Act, 1976, has been a major incentive to increased interest in the improvement of trout fisheries. 4. Brown trout stocks have been reduced in certain areas due to the effects of afforestation, acidification, land drainage and farm wastes. Various remedial measures have been proposed and implemented. 5. To meet the increasing demands for trout fishing, many loch and reservoir fisheries are now stocked with rainbow trout in preference to brown trout. Attention should be paid to the interaction of these two species in both standing and in running waters., where fish farm escapees and inadvisable releases go unrecorded. 7. Research into the genetic effects on wild stocks from the liberation of large numbers of hatchery-reared brown trout has been lacking and probably many ‘pure’ indigenous stocks have been lost. More work in this field is essential. 8. Proposals are outlined for future brown trout research and recommendations are made for better management. Suggestions are also put forward for changes in the legislation to further protect Scottish brown trout stocks.     

Assessing the status of British wild brown trout, Salmo trutta, stocks: a pilot study utilizing data from game fisheries

SUMMARY. 1. This paper describes an analysis of results from a questionnaire (Giles, 1987) which requested information on the status of game fish stocks (especially wild brown trout Salmo trutta L.) from those Game Conservancy members who own or manage fisheries. Also included are data arising from fishery log book records which are used to indicate trends in brown trout catch per unit effort (CPUE) over past decades.
2. The key finding from questionnaire returns was the widespread reporting of apparent declines of wild brown trout stocks in a total of twenty-seven sites throughout Britain. The affected waters are geographically widespread from the south-west of England through southern and south-east England, south Wales, northern England and Scotland including the Western Isles. Suspected stock declines of migratory (sea) trout, salmon Salmo salar L. and grayling Thymallus thymallus L. were also reported. Where brown trout occur alone, a significantly higher proportion of questionnaire returns recorded a stock decline compared with fisheries containing both brown and rainbow trout Salmo gairdneri Richardson.
3. Trout catch records from a small selection of differing fishery types are presented and the value of such data in fish stock assessment is discussed.     

Growth‐rate variation in brown trout in small neighbouring streams: evidence for density‐dependence?

The within- and among-population variation in individual growth rate of brown trout Salmo trutta L. was studied in five small neighbouring streams (seven isolated populations) within a distance of 70 km in east Norway. Observed growth rate was only weakly correlated with predicted maximum growth rate based on laboratory models, and there was a significant interaction with site. A generalized linear model showed that growth rate was positively correlated with temperature, but also that growth rate decreased as the summer season progressed. This might indicate either a seasonal decline in food availability or appetite, or a change in energy allocation strategy. Growth rate decreased with increasing fish age, probably as an effect of sexual maturation of older fish, and differential allocation of protein and lipid among different size-groups of brown trout. After adjusting for variation in temperature, season, and fish mass, there was still significant among-site variation in growth rate. A significant part of this variation was due to variation in brown trout density and the presence or absence of Alpine bullhead Cottus poecilopus . Growth rate decreased with increasing brown trout density, and was lower in the presence than in the absence of Alpine bullhead after correcting for variation in brown trout density. This last result may indicate the presence of interspecific competition.     

Inappropriate analysis does not reveal the ecological causes of evolution of stickleback armour: a critique of Spence et al. 2013

In a recent paper in this journal, Spence et al. (2013) sought to identify the ecological causes of morphological evolution in three‐spined sticklebacks Gasterosteus aculeatus, by examining phenotypic and environmental variation between populations on the island of North Uist, Scotland. However, by using simple qualitative assessments of phenotype and inappropriate measures of environmental variation, Spence et al. have come to a conclusion that is diametrically opposite to that which we have arrived at in studying the same populations. Our criticisms of their paper are threefold: (1) using a binomial qualitative measure of the variation in stickleback armour (“low” versus “minimal” (i.e., “normal” low‐plated freshwater sticklebacks versus spineless and/or plateless fish)) does not represent the full range of phenotypes that can be described by quantitative measures of the individual elements of armour. (2) Their use of unspecified test kits, with a probable accuracy of 4 ppm, may not be accurate in the range of water chemistry on North Uist (1 to 30 ppm calcium). (3) Their qualitative assessment of the abundance of brown trout Salmo trutta as the major predator of sticklebacks does not accurately describe the variation in brown trout abundance that is revealed by catch‐per‐unit‐effort statistics. Repeating Spence et al.'s analysis using our own measurements, we find, in direct contradiction to them, that variation in stickleback bony armour is strongly correlated with variation in trout abundance, and unrelated to variation in the concentration of calcium in the lochs in which they live. Field studies in ecology and evolution seldom address the same question in the same system at the same time, and it is salutary that in this rare instance two such studies arrived at diametrically opposite answers.     

The fish populations in inshore waters of the West Coast of Scotland. The distribution, abundance and growth of the whiting (Merlangius merlangus L.)

The whiting ( Merlangius merlangus ) is an important commercial species from the West Coast of Scotland. This paper describes the distribution, abundance, age composition and growth of juvenile whiting which are found in the inshore areas and sea lochs. No spawning was observed in the areas and it is assumed there was an active migration of whiting from the open sea. They were first caught by the bottom trawl in July, reached maximum abundance in November and December and thereafter declined in numbers until few remain by early summer. An exception to this occurs in Loch Etive and possible explanations for the fact that whiting tend to remain for longer periods and grow at a slower rate in this loch are discussed.     

Ice-cover effects on competitive interactions between two fish species

1. Variations in the strength of ecological interactions between seasons have received little attention, despite an increased focus on climate alterations on ecosystems. Particularly, the winter situation is often neglected when studying competitive interactions. In northern temperate freshwaters, winter implies low temperatures and reduced food availability, but also strong reduction in ambient light because of ice and snow cover. Here, we study how brown trout [Salmo trutta (L.)] respond to variations in ice-cover duration and competition with Arctic charr [Salvelinus alpinus (L.)], by linking laboratory-derived physiological performance and field data on variation in abundance among and within natural brown trout populations. 2. Both Arctic charr and brown trout reduced resting metabolic rate under simulated ice-cover (darkness) in the laboratory, compared to no ice (6-h daylight). However, in contrast to brown trout, Arctic charr was able to obtain positive growth rate in darkness and had higher food intake in tank experiments than brown trout. Arctic charr also performed better (lower energy loss) under simulated ice-cover in a semi-natural environment with natural food supply. 3. When comparing brown trout biomass across 190 Norwegian lakes along a climate gradient, longer ice-covered duration decreased the biomass only in lakes where brown trout lived together with Arctic charr. We were not able to detect any effect of ice-cover on brown trout biomass in lakes where brown trout was the only fish species. 4. Similarly, a 25-year time series from a lake with both brown trout and Arctic charr showed that brown trout population growth rate depended on the interaction between ice breakup date and Arctic charr abundance. High charr abundance was correlated with low trout population growth rate only in combination with long winters. 5. In conclusion, the two species differed in performance under ice, and the observed outcome of competition in natural populations was strongly dependent on duration of the ice-covered period. Our study shows that changes in ice phenology may alter species interactions in Northern aquatic systems. Increased knowledge of how adaptations to winter conditions differ among coexisting species is therefore vital for our understanding of ecological impacts of climate change.     

Reproductive isolation and genetic differentiation of ferox trout from sympatric brown trout in Loch Awe and Loch Laggan, Scotland

Molecular marker studies reported here, involving allozymes, mitochondrial DNA and microsatellites, demonstrate that ferox brown trout Salmo trutta in Lochs Awe and Laggan, Scotland, are reproductively isolated and genetically distinct from co-occurring brown trout. Ferox were shown to spawn primarily, and possibly solely, in a single large river in each lake system making them particularly vulnerable to environmental changes. Although a low level of introgression seems to have occurred with sympatric brown trout, possibly as a result of human-induced habitat alterations and stocking, ferox trout in these two lakes meet the requirements for classification as a distinct biological, phylogenetic and morphological species. It is proposed that the scientific name Salmo ferox Jardine, 1835 , as already applied to Lough Melvin (Ireland) ferox, should be extended to Awe and Laggan ferox.     

Sea trout in North Argyll sea lochs: II. diet

The diet of sea trout in some of the sea lochs of the west coast of Scotland was investigated. The contents of 986 sea trout stomachs from the Loch Etive area (1970–1973), and 291 stomachs from the Loch Eil area (1964–1973), were examined and the composition of the diet, seasonal changes and the effect of trout size were analysed using frequency of occurrence, dry weight and number of organisms methods. Benthic feeding (crustacea and annelids) was more important in winter while midwater and surface organisms (young fish and insects) were preferred in summer. Young fish (mainly clupeids and sand eels) featured more in the diet of larger trout (≤21 cm) than in the smaller size range (≥21 cm). Surveys of 24-h were completed in June and September, 1972, and indicated that availability of food was the main factor influencing the presence or absence of trout. Bottom feeding was greatest during the day while the amount of midwater and surface feeding tended to increase between sunset and sunrise.     

The fresh waters of Shetland: Physical and morphometric characteristics of lochs

SUMMARY. 1. Basic physical and morphometric data are presented for sixty-five lochs on the Shetland Islands. The interrelations between loch dimensions are comparable to those recorded elsewhere in Scotland, but the smaller lochs on Shetland tend to be somewhat shallower, and the large lochs somewhat deeper than usual.
2. The results of a multivariate analysis show that most of the physical variables measured in the field are closely correlated with easily measured map attributes.     

Partial migration in a landlocked brown trout population

Population densities of landlocked lake‐migratory brown trout Salmo trutta were estimated in two distinct lotic sections, separated by a lentic segment, in the Greåna River, Sweden, and individual growth and habitat use were monitored for 835 tagged brown trout from September 1998 to June 2000. Residency dominated in the upstream section where density of 0+ and 1+ year brown trout was low and growth rate high. In contrast, >90% of the brown trout that migrated to the lake originated from the downstream section, where density was high and growth rate low. For ≥2+ year individuals, growth rate was similar between the two stream sections, but densities were higher in the upstream than in the downstream section. Lake‐migrants had higher growth rates than non‐migrants (residents) during the autumn of both years. From September to May, migrants increased their body mass by >35%, whereas non‐migrants increased by <5%. Approximately 70% of the brown trout moved <10 m and <2% moved between the two stream sections, indicating that the lentic habitat might function as a barrier for juveniles. Differences in migratory behaviour, density and growth between the upstream and the downstream section might indicate that environmental factors influence the decision to migrate. It cannot be excluded, however, that the observed differences are genetically programmed, selected by migration costs that favour migratory behaviour downstream and residency upstream.     

Recapture rate and growth of hatchery‐reared brown trout (Salmo trutta v. fario,L.) in Blanice River and the effect of stocking on wild brown trout and grayling (Thymallus thymallus,L.)

Hatchery‐reared adult brown trout, Salmo trutta v. fario L., [215–335 mm standard length (L_S), n = 82] were individually tagged and released into three sections of the Blanice River in May 2007. Wild populations of brown trout and grayling, Thymallus thymallus, L., in these sections and three non‐stocked control sections were also tagged. The recapture rate of hatchery‐reared adult brown trout after 6 months (18%, n = 15) was comparable to that of wild adult brown trout in stocked (15%, n = 14) and control (14%, n = 11) sections. The recapture rates of wild brown trout and grayling after 6 months were higher in control sections than in stocked sections, but the differences were not significant. The movement of recaptured large juvenile wild brown trout from stocked sections was significantly higher (36%) than from control sections (9%). Wild brown trout growth and grayling growth were unaffected by stocking with adult hatchery‐reared brown trout.     

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.     

Landscape factors that shape a slow and persistent aquatic invasion: brown trout in Newfoundland 1883–2010

Aim We investigated watershed‐scale abiotic environmental factors associated with population establishment of one of the ‘world’s 100 worst alien invaders’ on a temperate Atlantic island. Within the context of the conservation implications, we aimed to quantify (1) the early history and demographics (numbers and origins) of human‐mediated brown trout (Salmo trutta) introductions, (2) the current distribution of established populations, and (3) the watershed‐scale environmental factors that may resist or facilitate trout establishment. Location Island of Newfoundland, Canada. Methods We combined field sampling with historical and contemporary records from literature to assemble a presence–absence and physical habitat database for 312 watersheds on Newfoundland. Probability of watershed establishment was modelled with general additive ANCOVA models to control for nonlinear effects of propagule pressure (i.e. the distance to and number of invasion foci within a biologically relevant range) and model performance based on AIC. Results Between 1883 and 1906, 16 watersheds were introduced with brown trout from the Howietoun Hatchery, near Stirling, Scotland. Since that time, populations have established in 51 additional watersheds at an estimated rate of spread of 4 km per year. We did not detect any obvious abiotic barriers to resist trout establishment, but showed that for a given amount of propagule pressure that relatively large and productive watersheds were most likely to be established. Main conclusions Brown trout have successfully invaded and established populations in watersheds of Newfoundland and are currently slowly expanding on the island. Populations are more likely to establish in relatively large and productive watersheds, thereby supporting predictions of island biogeography theory. However, we suggest that all watersheds in Newfoundland are potentially susceptible to successful brown trout invasion and that abiotic factors alone are unlikely to act sufficiently as barriers to population establishment.     

The status of brown and rainbow trout, Salmo trutta and S. gairdneri as hosts of the acanthocephalan, Pomphorhynchus laevis

The status of brown and rainbow trout as hosts of Pomphorhynchus laevis was studied in the field and by means of laboratory investigations. Field data indicated that rainbow trout might belong to the group of preferred hosts of P. laevis , whereas brown trout belonged to the group in which the parasite achieved less than optimal growth and maturation. This was confirmed by laboratory infections. In rainbow trout P. laevis attained up to three times the growth rate in brown trout and maturation occurred whereas in brown trout establishment was lower, growth slower and no parasites matured. Changes in the behaviour of infected Gammarus pulex induced by the presence of P. laevis cystacanths were such as to render the shrimps more vulnerable to predation by trout and other surface and mid-water feeding fish, and selective predation upon infeged G. pulex by fish was demonstrated. nvestigations into the stimuli necessary for eversion of cystacanths of P. laevis revealed that the most important factor was a non-specific component of bile, and it was concluded that cystacanths were likely to evert in any species of fish. Recognition of the different status of brown and rainbow trout as hosts of P. laevis still fails to explain some peculiarities in the distribution of the parasite in the British Isles, where in Britain it occurs in trout in only one river but in Ireland in all rivers throughout the country. It is suggested that the Irish parasites may constitute a different strain of P. laevis , since they use a different species of intermediate host and are better able to survive in brown trout.     

A Bioenergetic Analysis of Factors Limiting Brown Trout Growth in an Ozark Tailwater River

We examined prey utilization and energy consumption by brown trout, Salmo trutta, in a cold tailwater (Little Red River, Arkansas, USA; LRR) having low biodiversity and low availability of fish as prey. Stomach content analysis and age estimation were performed on thirty brown trout (10 each of three size classes for a total of 710 trout) collected monthly from an upstream and downstream site over a 1-year period. Diet diversity was low at both sites, as 80% and 70% of all prey consumed by upstream and downstream brown trout, respectively, were isopods. Piscivory (<0.5% of individuals sampled) and consumption of terrestrial invertebrates were rare. There was no relation between diet diversity and trout age, and a very small ontogenetic shift in brown trout diet. Second, we investigated brown trout growth rates relative to prey consumption and temperature. Temperatures and availability of prey were less than required for maximal trout growth. However, prey availability limited trout growth directly, but sub-optimal temperatures probably buffered the effect of this reduced energy consumption by reducing metabolic energy expenditures. Brown trout growth was 54.8–57.0% of the maximum predicted by a bioenergetics model. Instantaneous growth rates for age 1 and adult brown trout were slightly higher for those downstream (0.195) versus those upstream (0.152). Although isopods are abundant within this tailwater to serve as a forage base, the displacement of native fish fauna and subsequent lack of establishment of cold-tolerant forage fish species due to the thermal regime of hypolimnetic release from Greers Ferry Reservoir probably serves as a major barrier to brown trout growth.     

An aeromonad epidemic in the brown trout (Salmo trutta L.)

The pathology and bacteriology of an aeromonad epidemic in the spawning population of brown trout ( Salmo trutta L.) of the freshwater Loch Leven, Kinross, Scotland, is described, together with estimates of its quantitative effect on the stock of trout.     

Invasive brown trout Salmo trutta induce differential growth strategies in the native snow trout Schizothorax richardsonii of Himalaya: Are natives in unaltered rivers better at picking the gauntlet of invasion?

Brown trout Salmo trutta is a potent global invader and its establishments have progressively altered physiologies, life-histories and niche-availabilities for native fish species. River impoundments further escalate its invasion potential. The Himalayan rivers however, stay uncharted for the effects of brown trout interactions with the native fish fauna. Snow trout Schizothorax richardsonii a Himalayan cold-water native, concerningly overlaps its range with brown trout. To understand its responses to invasion pressures, we investigated brown trout effects on the age and growth of snow trout populations in three rivers with varying levels of perturbation: (a) a dammed and (b) an undammed river with the invasive brown trout in comparison to (c) an undammed river without invasion pressures. We found sympatric snow trout in the undammed river to respond to brown trout invasion with fast life history responses, showing an early age-at-maturity (A₅₀ = 1.2 years) and fast growth with a higher growth constant (K = 0.40 yr⁻¹) and specific rate of linear growth across life. On the contrary, sympatric snow trout in the dammed river showed an explicitly slow life-history by maturing at a higher age (A₅₀ = 2.9 years) and a slow growth, with a lower growth constant (K = 0.26 yr⁻¹) and specific linear growth rates. Our findings suggest that, the snow trout appear to present stronger response to brown trout invasions when the river is unaltered and free from hydropower operations and damming. Further research is strongly warranted from other high-altitude Himalayan basins to delineate the variation in growth strategies exhibited by snow trout in sympatry with the invasives.