Environmentally induced spatio-temporal variations in the fecundity of brown trout Salmo trutta L.: trade-offs between egg size and number

1. Resident brown trout Salmo trutta in the Esva River basin (north Spain) live in a patchy environment with tracts of riparian forest or meadow along stream banks. This study assessed whether the reproductive traits of brown trout from four contrasting sites reflected site-specific factors.
2. Length at maturity (10.5–11 cm of 1 + individuals) was the same in the four sites examined but slowest growers in slow-growing sub-populations delayed maturity for 1 year relative to fast-growing fish. The analysis of monthly variations in egg size and number suggest that two 'decisions' in two consecutive years are required to complete spawning. The first concerns the number of eggs, determined when trout are still 0 +, and the second concerns egg size.
3. At three sites, egg size and number did not differ significantly between years but highly significant interannual variations were apparent at another site. Fish length was the major determinant of egg size and number at all sites but for any given length, brown trout at sites where the fish exhibited higher growth rates spawned more, but smaller, eggs than those at slow-growing sites. This spatial pattern was identical to the temporal pattern exhibited by trout at another site. The combination of temporal (year-to-year) and spatial (between rivers) variations in egg size and number showed a significant negative correlation, supporting the operation of a trade-off between these two traits.
4. The trade-off between egg size and number seems to be determined by site-specific factors, with slow-growing trout at sites which are fully covered by canopy spawning fewer, but larger, eggs than fast-growers in unshaded sites.     

Growth, production and bioenergetics of brown trout in upland streams with contrasting riparian vegetation

1. A series of laboratory-based equations on trout growth and bioenergetics developed by J.M. Elliott were applied to data collected for brown trout ( Salmo trutta L.) under field conditions in Co. Mayo, Western Ireland. Fish were collected by electrofishing eight upland streams with contrasting riparian vegetation; grassland, open canopy and closed canopy deciduous.
2. Stream temperatures, one of the main influencing factors on fish growth and energetics, did not differ significantly between riparian types.
3. Observed growth rates were lower than the predicted maximum growth rates and were not influenced by riparian vegetation type. Growth ranged between 0.66% day⁻¹ for 0 + trout to 0.08% day⁻¹ for 2 + trout.
4. Production estimates showed no clear difference between riparian vegetation types over the growing season.
5. Fish densities and biomass tended to be greater in closed canopy streams particularly in summer.
6. Actual ration sizes calculated for trout were similar to the ration required for maintenance metabolism and were only 45–63% of the maximum potential rations. Although there was an ontogenetic increase in ration size with increasing fish age, the proportion of ration available for growth (i.e. the difference between actual and maintenance rations) did not differ between age classes but was greatest in summer. 1+ and 2+ trout show greatest ration available for growth in grassland streams.
7. Trout growth did not differ between riparian vegetation types but did vary seasonally with greatest attainment in summer. Growth was limited in the present study possibly due to combined effects of reduced prey available to fish and low stream temperatures reducing metabolic requirements. In such food limited systems, terrestrial invertebrate energy subsidies could have significant benefits to brown trout growth, production and bioenergetics.     

Environmental factors and life histories of isolated river stocks of brown trout (Salmo trutta m. fario) in Søre Osa river system,Norway

Synopsis The brown trout in the Søre Osa river system are isolated from the lake above by a dam. They are generally of a small size, but their condition coefficient is high. They reach sexual maturity at an early age and have a short life span. The substratum in the Søre Osa mainly consists of large stones covered with a dense mossy vegetation that creates good fish cover. A high biomass of zoobenthos gives an adequate food supply throughout the year. The trout spawn mainly in tributaries, where the water level is unpredictable, some years being too low to permit successful spawning for the migrating individuals in the stock. The life history of these fish depends on the amount of food available in the habitat and on reproduction in unstable environments where density-independent mortality factors appear important. Many young migrate to the main stream during their first year. In the tributaries, there is an excess of resident males and it is hypothesized that a population structure with small resident males and large migratory males is maintained by partial inbreeding and kin selection. In the main stream the trout are larger and they have higher growth rates in the upper part than farther downstream. This is probably so because the food particles that are flushed down from the lake above provide a better supply of zoobenthic food.     

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.     

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

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

Movements of brown trout, Salmo trutta, and juvenile Atlantic salmon, Salmo salar, in a coastal stream in northern Norway

Movements of resident brown trout (age 2+ to 9+ years) and young Atlantic salmon (age 1+), stocked as fry, were studied in July, August and September in a coastal stream in northern Norway. Between 85 and 89% of the brown trout were recaptured in the study area (346m, 1326m²) within 45m of their release point throughout the investigation period. Most specimens had moved less than 150m. Trout movements were related to local variation in density. Trout occupying those sections of stream with the lowest fish densities (5.3–10.9 fish 100m^?2) had a significantly lower movement rate than fish from sections with densities between 13.7 and 31.5 fish 100m^?2. Trout that moved from their marking section were significantly larger than specimens that did not leave their original site. There was a significant correlation between permanence of station each month and the mean water level that month. The majority of the trout (47%) were caught at undercut stream banks or at sites in the proximity of these. A decrease in water level during the study period resulted in a high loss (36%) of such habitat, probably forcing some individuals to move. The recapture data indicate that the trout population consists of one small (c. 15–20%) mobile, and one large sedentary component. Young salmon displayed high station permanence in July and August (93% and 96%). However, in the autumn they exhibited a significant downstream movement, and only 73% were recaptured within their release section. This movement was significantly higher for larger specimens, and is thought to occur because of a pre-winter change in habitat, initiated by a decline in stream temperature. In contrast to trout, salmon in sections containing the lowest densities (22.0–25.0 fish 100m^?2) did not show significantly lower movement rates when compared with salmon at higher densities (32.2–46.3 and 51.8–60.6 fish 100m^?2). The spatial distribution of young salmon indicated the formation of territorial mosaics over the stream bed, which are thought to reduce intraspecific competition.     

The ecology of brown trout Salmo trutta in English chalk streams

SUMMARY. .1. Chalkstream trout are fast-growing, short-lived with a stable age structure that is a consequence of the lack of extremes in the abiotic environment, especially of water temperature and flow.
2. Levels of secondary (invertebrate) production arc high and there is no evidence that interspecific or intraspecific competition for food resources limits growth.
3. Dietary studies show a broad similarity between trout and other species, but indicate differences that reflect variations in the micro-habitat distributions between fish species.
4. Trout numbers appear to be limited initially by the availability of gravel spawning areas, and then by areas suitable for newly-emerged fry. Reduction in stream discharge in the spring, either naturally or by man, can lower the number of 0+ trout that survive.
5. Eels are not important predators on trout eggs or fry. but a reduction in pike numbers can lead to a decrease in the mean weight of pike. Small pike do not pose a serious threat to stocked yearling trout.     

Habitat utilization by sympatric arctic charr Salvelinus alpinus L. and brown trout Salmo trutta L. in Lake Atnsjø, south-east Norway

SUMMARY. 1. Habitat utilization, as well as inter- and intraspecific relations of different size groups of arctic charr (Salvelinus alpinus (L.)) and brown trout (Salmo trutta L.) in Lake Atnsjø, south-east Norway, were investigated by analysing food and spatial niches from monthly benthic and pelagic gillnet catches during June-October 1985.
2. Small individuals (150–230 mm) of both arctic charr and brown trout occurred in shallow benthic habitats. However, they were spatially segregated as arctic charr dominated at depths of 5–15 m and brown trout at depths of 0–5 m.
3. Larger (>230 mm) arctic charr and brown trout coexisted in the pelagic zone. Both species occurred mainly in the uppermost 2-3 m of the pelagic, except in August, when arctic charr occurred at high densities throughout the 0–12 m depth interval. On this occasion, arctic charr were segregated in depth according to size, with significantly larger fish in the top 6 m. This was probably due to increased intraspecific competition for food.
4. The two species differed in food choice in both habitats, Arctic charr fed almost exclusively on zooplankton, whereas brown trout had a more variable diet, consisting of surface insects, zooplankton. aquatic insects and fish.
5. The data suggest that the uppermost pelagic was the more favourable habitat for both species. Large individuals having high social position occupied this habitat, whereas small individuals lived in benthic habitat where they were less vulnerable to agonistic behaviour from larger individuals and less exposed to predators. The more aggressive and dominant brown trout occupied the more rewarding part of the benthic habitat.     

Stocking hatchery‐reared brown trout in different densities into a wild population – a comparison of growth and movement

In spring 2001 and 2002 a small stream was stocked with tagged hatchery‐reared yearling brown trout ( Salmo trutta ), in order to study their influence on the resident brown trout population. The stream was separated into six sections: two sections without stocking, two sections where stocking doubled the trout population and two sections where the fish population was quadrupled. The working hypothesis was that due to food limitation (competition) growth of the wild fish will be negatively influenced by stocking, and wild fish will be displaced by the (possibly more aggressive) hatchery fish. Surprisingly, growth rate of wild and stocked fish of the same age was similar and independent of stocking density. Two main reasons may be responsible for this finding: only a low percentage of the stocked fish remained in the stream, and food was not limited during summer. Only 12–19% of the stocked fish were recaptured after six months, in contrats to 40–70% of one‐year old and up to 100% of older wild trout. The wild fish were not displaced by hatchery‐reared fish: During summer the wild fish remained more or less stationary, whereas most of the stocked trout had left their release site. The results indicate that in a natural stream stocking of hatchery reared brown trout does not influence negatively growth and movement of the wild fish independent of stocking density.     

Life history and habitat use of Norwegian brown trout (Salmo trutta)

SUMMARY. 1. Brown trout ( Salmo trutta) life history and habitat use were studied in two Norwegian rivers: the Vosso river system, western Norway, and the Søre Osa, eastern Norway.
2. Age-groups were partly segregated in feeding habitats, the youngest fish living mainly in running water and in the littoral zone of lakes, the older fish also exploiting pelagic waters and deeper epibenthic habitats. In a population with free access to and from the sea, some individuals smoltified and became sea-run migrants, performing yearly migrations to the coastal sea, whereas others stayed as freshwater residents throughout their entire life span.
3. Within local populations, females were larger and less variable in size than males. This was partly because females matured at an older age than males, partly because the sexes tended to exploit feeding habitats with different food and growth Conditions. Within age-groups, females were more pelagic and migrated more than males, whereas males were more confined to running water and epibenthic areas than females. In the pelagic zone, males were more abundant in near-surface water, and females more abundant in deeper areas. When exploiting the same feeding areas, the two sexes grew at the same rate. There therefore appears to be a connection between feeding habitat and the reproductive ecology of brown trout.     

Ecological Forms of Black Sea Brown Trout (<Emphasis Type="Italic">Salmo trutta labrax</Emphasis>) in the Mzymta River as Manifestation of Ontogenetic Plasticity

Populations of brown trout in the Mzymta River and its tributaries include anadromous (mainly female) and resident (mainly males) fish. Some resident males in the basin of the Mzymty River attain sexual maturity at the age 1+, and resident females mature at the age 2+ or 3+. The maximum age of resident fish is 4+ in the samples studied. Migrations of anadromous brown trout to the sea occur at the ages 1+, 2+, or 3+. Future spawners spend from 1 to 4 years at feeding grounds in the sea. Smolts of the population are characterized by performing not only spring but also autumn migrations to the sea. One smolt specimen has been detected upstream from the dam in the river where spawners of anadromous brown trout do not migrate; this means that the capability for sea migrations persists long in the population represented only by resident specimens of brown trout. The diversity of life cycles and ecological forms in populations of brown trout is not lower than in populations of brown trout in Northern and Western Europe. The comparison of the data obtained with published data makes it possible to come to the conclusion about the high plasticity of ontogenesis of Black Sea brown trout.     

Home range of juvenile Atlantic salmon, Salmo salar, and brown trout, Salmo trutta, in a Norwegian stream

SUMMARY. 1. The sizes of home ranges of juvenile Atlantic salmon (age 1 +) and brown trout (age 2+ to 9+) in a Norwegian coastal stream were estimated by local movements of batch-marked fish from 12.5 and 25 m long sections. Only recoveries made in the release section and in up-and downstream neighbouring sections were considered.
2. There was no significant difference in the average percentage of recaptures of salmon and trout between 12.5 and 25 m sections; a stream area of about 40–50 m² defines the size of home range for stocks of both species.
3. The fraction of brown trout recaptured in release sections increased with increasing fish densities, indicating a smaller home range under these conditions.     

Density reductions by predatory trout increase adult size and fecundity of surviving caddisfly larvae in a detritus-based stream food web

1. In some situations, individuals surviving in environments where predation is intense can grow faster because the benefits of release from intraspecific competition outweigh costs associated with anti-predator responses. Whether these 'thinning' effects of predation occur in detritus-based food webs where resource renewal occurs independently of consumption by consumers was studied. We investigated how effects of predatory brown trout ( Salmo trutta ) on the larvae of the detritivorous stream caddisfly, Zelandopsyche ingens , influenced the size and fecundity of the caddisfly adults.
2. Trout substantially reduced the abundance of Z. ingens larvae, but adult male and female Z. ingens were significantly larger in trout streams compared to fishless streams. Females in trout streams had 33% more eggs than fishless stream females, and egg sizes were not significantly different. In mesocosms, Z. ingens larvae in low density treatments reflecting trout stream abundances grew significantly faster than larvae in high density treatments that were characteristic of fishless stream abundances. Non-lethal trout presence did not influence case building behaviour, feeding rates or growth or Z. ingens larvae, indicating non-lethal effects of predators were negligible.
3. Increased adult size and fecundity associated with trout stream individuals were probably a result of predator thinning of larval density indirectly releasing surviving Z. ingens from intraspecific competition. Thus, predator thinning did influence interactions between larvae in this detritus-based food web as larval growth was strongly density-dependent. However, extrapolating the total number of eggs potentially produced indicates the increased fecundity of females in trout streams would not compensate for losses of larvae to trout predation.     

Movement and site fidelity in young brown trout Salmo trutta populations in a southern Irish stream

Populations of brown trout Salmo trutta were monitored at a number of sites within a single stream, using an individual marking technique and recapturing uniquely marked fish repeatedly over a period of 12 months. Individual 1 + and 2 + resident brown trout in the Glenfinish River were found to consist of stationary and mobile component populations. The latter population consisted of a number of individuals observed moving mostly in an upstream direction, within a range of 0.03–2.24 km. On a large spatial scale, individuals in the stationary component population exhibited some degree of home site fidelity within the stream, over a period of 3–4 months, after which the fish tended to move from the site. Within sites, fidelity to either riffle or pool habitats, mostly the latter, was apparent in a proportion of the population. On a smaller scale, fidelity to the exact position with respect to boulders in the stream was also evident in a number of individuals. Home range size was calculated amongst these individuals, with ranges of up to 20 m recorded.     

Reproductive traits of stream-dwelling brown trout Salmo trutta in contrasting neighbouring rivers of central Spain

1. Reproductive traits were studied in seven wild populations of resident brown trout in Spain. We examined whether growth, and certain environmental conditions such as water temperature or food abundance, could explain interpopulation variation in the reproductive characters.
2. The results indicated that nearby populations subjected to a wide variation in environmental conditions exhibited a similar reproductive performance.
3. Age distributions and sex ratios were not significantly different among populations. Age ranged from 0+ to 4+ years but the populations were dominated by the 0+ to 2+ groups. In all rivers females matured at age 2+ as opposed to age 1+ and 2+ for males. Mean length at the end of the growth period differed significantly among populations. However, annual growth rate was similar among rivers and was not significantly correlated with either biomass of the benthos or water temperature during the growth season.
4. Reproductive effort, body condition, fecundity and egg size did not correlate significantly with either the abundance of benthic invertebrates or water temperature. Once the effect of body length on both egg size and number was removed, a significant negative correlation was found between these two traits among populations.     

Pre‐migratory differentiation of wild brown trout into migrant and resident individuals

In February to March, wild brown trout Salmo trutta were captured by electrofishing in a natural watercourse (tributaries of the River Lille Aa, Denmark), individually tagged (Passive Integrated Transponders), and released. Representatives of the tagged brown trout were recaptured on the release sites in April by electrofishing and eventually caught in downstream smolt traps ('migrants') placed in the main river or by electrofishing ('residents') on the initial sites in June. Upon each capture, smolt appearance and body size were evaluated, and a non‐lethal gill biopsy was taken and used for Na⁺,K⁺‐ATPase analysis. Based on repetitive gill enzyme analysis in individual fish, a retrospective analysis of the rate of development in individual brown trout ultimately classified as migrants or residents was performed. Two months prior to migration, a bimodal morphological and physiological (gill Na⁺,K⁺‐ATPase) development concurred and was related to the subsequent differentiation into resident and migratory fractions of each population. This differentiation was unrelated to growth rate and body size of individual fish but skewed in favour of migratory females. Individuals destined to become migrants developed a smolt‐like appearance before the onset of migration and had higher rate of change of gill Na⁺,K⁺‐ATPase activity than fish remaining residents. The rate of change of gill Na⁺,K⁺‐ATPase activity was independent of the distance migrated to the trap (3–28 km). Thus in bimodal wild brown trout populations a major increase in enzyme activity takes place before migration is initiated and is a characteristic of migratory individuals only.     

Interacting effects of temperature and density on individual growth performance in a wild population of brown trout

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Use of space and food by resident and migrant brown trout,Salmo trutta

Synopsis Parr and resident forms of brown trout,Salmo trutta, from Vangsvatnet Lake, Norway live in freshwater, while migrant forms live in coastal waters during summer and in freshwater during winter. About 80% of parr and residents live at depths <5 m, smolts and migrants are more confined to near-surface water. Brown trout partly segregate by size, age and sex from spring through autumn. More than 90% of parr and residents in the tributaries are 0–2 years old, 2–14 cm in length, in the littoral zone 0–3 years old, 7–24 cm in length, and in the pelagic zone 2–6 years old, 18–32 cm in length. The mean body length of equal-aged fish increases from tributaries through littoral to pelagic zones in the lake. Smolts (2–7 years, 14–29 cm) leave the lake from April through August and return during September–October. Migrants (2–11 years, 23–67 cm) leave the take in April–May and return during August–September; sexually mature fish return earlier than immatures. Female brown trout are less stream-dwelling, but more migrant and pelagic than males. Most individuals in the lake population spend the winter in the littoral zone. In the tributaries, diet differs significantly between age-groups of parr; young fish feed on smaller food items than do older fish. In the lake, parr and residents living in the same habitats feed on the same food items. Littoral brown trout feed mainly on insect larvae and chironomid pupae, pelagic brown trout feed on zooplankton and surface insects. Migrants feed little while staying in freshwater, except for matures which feed on young salmonids and surface arthropods during the 2 first months after they had returned from coastal waters. The results are discussed in relation to growth possibilities and mortality risks of the different habitats.Reprint request to B. Jonsson.     

Fitness-related effects of growth investment in brown trout under field and hatchery conditions

The mortality of brown trout Salmo trutta over winter in a near-natural stream was not significantly increased by growth hormone (GH) treatment, but lipid reserves were lower in GH-treated fish. As GH-treated trout grew faster than controls, GH appears to promote growth at the expense of investment in maintenance. However, the growth promoting effect of GH was much more pronounced in the hatchery than in the stream, suggesting that the pay-off associated with increased growth investment is higher under hatchery conditions with unrestricted food supply than in wild, where food availability is limited.     

Seasonal and regional infestation characteristics of three ectoparasites of whiting, Merlangim merlangus L., in the North Sea

The colonization by both resident and migrating spawner populations of brown trout and the characteristics of resident and migrating juveniles derived from the two populations have been studied in a brook and its tributary over 4 years. Resident trout spawns mainly in the upstream part of the brook and migrating trout in the downstream part. There are density and growth variations for the two age classes (0+ and 1 +) of juveniles in autumn according to the year and the environment. In the brook, the population of 0 + fish increases from downstream to upstream while the density of other age classes decreases. The migrating juvenile population of the brook changes annually and consists mainly of 1 s (one summer) individuals coming from the upper part. These individuals migrate generally in autumn and winter while young trout produced in the middle and downstream parts of the brook migrate mainly in the spring. The emigration process of the 0 + population decreases markedly from upstream to downstream and appears to be independent of the autumn length and sex ratio. In the tributary, most trout are 0+ years old, the population structure is different, and no migrating fish is observed. The results are discussed and a colonization strategy of the brown trout population in this brook is suggested.