Effects of body size on sympatric shelter use in over‐wintering juvenile salmonids

The study tests two hypotheses: (1) the degree of shelter dominance in Atlantic salmon Salmo salar and brown trout Salmo trutta increases progressively with increasing size differential between heterospecific fish in a pair and (2) shelter dominance, standardized to size differential, correlates with aggression. The results support the first but not the second hypothesis, suggesting that the fitness consequences of high growth performance during the summer are likely to become evident during winter. At this time of year, when mortality is high among both Atlantic salmon and brown trout, shelter dominance may increase the chances of survival.     

Behavioural strategies for species co‐existence during environmental extremes: competition between trout and salmon parr during dewatering

Competition between species of animals can be predicted to be intense under extreme environmental conditions during which niche overlap increases. Fluvial aquatic systems fluctuate naturally across a broad range of time‐scales. Land management activities impose further extreme fluctuations, particularly when water is abstracted for irrigation during summer. This study focused on the interaction between Atlantic salmon and brown trout parr during acute dewatering events. Brown trout are known to compete strongly for pool habitat whereas salmon can coexist by using riffle areas during normal flows. It is not known, however, how competition between the species affects their behavioural responses to extreme low flows when riffle areas decrease. Replicated groups of salmon were held in allopatry and sympatry with trout in sections of a large indoor stream, each of which was landscaped into riffles and pools. Space use and behaviours of the fish were recorded by direct visual observations and a network of Passive Integrated Transponder detectors. Here we report the response of the fish to dewatering and consider the system as a model for natural and forced responses of communities to environmental extremes.     

The evolution of cooperation and advanced social behaviour

Competition between species of animals can be predicted to be intense under extreme environmental conditions during which niche overlap increases. Fluvial aquatic systems fluctuate naturally across a broad range of time‐scales. Land management activities impose further extreme fluctuations, particularly when water is abstracted for irrigation during summer. This study focused on the interaction between Atlantic salmon and brown trout parr during acute dewatering events. Brown trout are known to compete strongly for pool habitat whereas salmon can coexist by using riffle areas during normal flows. It is not known, however, how competition between the species affects their behavioural responses to extreme low flows when riffle areas decrease. Replicated groups of salmon were held in allopatry and sympatry with trout in sections of a large indoor stream, each of which was landscaped into riffles and pools. Space use and behaviours of the fish were recorded by direct visual observations and a network of Passive Integrated Transponder detectors. Here we report the response of the fish to dewatering and consider the system as a model for natural and forced responses of communities to environmental extremes.     

Diet partitioning in sympatric Atlantic salmon and brown trout in streams with contrasting riparian vegetation

Prey intake by Atlantic salmon Salmo salar and brown trout Salmo trutta was measured across different riparian vegetation types: grassland, open canopy deciduous and closed canopy deciduous, in upland streams in County Mayo, Western Ireland. Fishes were collected by electrofishing while invertebrates were sampled from the benthos using a Surber sampler and drifting invertebrates collected in drift traps. Aquatic invertebrates dominated prey numbers in the diets of 0+ year Atlantic salmon and brown trout and 1+ year Atlantic salmon, whereas terrestrial invertebrates were of greater importance for diets of 1+ and 2+ year brown trout. Terrestrial prey biomass was generally greater than aquatic prey for 1+ and 2+ year brown trout across seasons and riparian types. Prey intake was greatest in spring and summer and least in autumn apart from 2+ year brown trout that sustained feeding into autumn. Total prey numbers captured tended to be greater for all age classes in streams with deciduous riparian canopy. Atlantic salmon consumed more aquatic prey and brown trout more terrestrial prey with an ontogenetic increase in prey species richness and diversity. Atlantic salmon and brown trout diets were most similar in summer. Terrestrial invertebrates provided an important energy subsidy particularly for brown trout. In grassland streams, each fish age class was strongly associated with aquatic, mainly benthic invertebrates. In streams with deciduous riparian canopy cover, diet composition partitioned between conspecifics with older brown trout associated with surface drifting terrestrial invertebrates and older Atlantic salmon associated with aquatic invertebrates with a high drift propensity in the water column and 0+ year fish feeding on benthic aquatic invertebrates. Deciduous riparian canopy cover may therefore facilitate vertical partitioning of feeding position within the water column between sympatric Atlantic salmon and brown trout. Implications for riparian management are discussed.     

Seasonal changes in the body composition of young riverine Atlantic salmon and brown trout

The body composition of protein and fat in Atlantic salmon Salmo salar and brown trout Salmo trutta before and after winter was investigated in a temperate, small river, normally ice covered from the middle of November until the end of March. Fat, protein and specific energy declined greatly in winter but were replenished rapidly in spring. Rates of decline were slower for the smallest fish, which also had the lowest specific content of fat, protein, and energy, while the differences in absolute amounts were greatest for the largest fish. The mean specific fat content was reduced by 45–70% during winter, relative to the pre-winter period (September). Mean daily reductions in specific enegy of the larger size groups of brown trout (3·7 × 10⁻³ kJ g⁻¹ day⁻¹) were almost half of the corresponding values for the largest Atlantic salmon (6·3 × 10⁻³ kJ g⁻¹ day⁻¹) during winter. A minor reduction in protein content was found during winter, with mean reductions of 6–10% in comparison to those in September. During spring the fat content was replenished rapidly, particularly for the smallest salmon fry (a threefold increase from April to June). Fat content in the larger salmon and trout increased by about 1·8 times. Based on estimated metabolic rates, digested energy during wintertime may contribute about two-thirds of the brown trout fry's energy demand. For Atlantic salmon, the corresponding value is about 50%. The winter period put considerable stress on the young salmonids living in lotic environments, in particular for the smallest fry with the lowest energy content before winter and the largest losses during winter. This should make the fry more vulnerable to adverse abiotic and biotic factors.     

Migration of hatchery‐reared Atlantic salmon and wild anadromous brown trout post‐smolts in a Norwegian fjord system

Hatchery‐reared Atlantic salmon Salmo salar ( n  = 25) and wild anadromous brown trout (sea trout) Salmo trutta ( n  = 15) smolts were tagged with coded acoustic transmitters and released at the mouth of the River Eira on the west coast of Norway. Data logging receivers recorded the fish during their outward migration at 9, 32, 48 and 77 km from the release site. Seventeen Atlantic salmon (68%) and eight sea trout (53%) were recorded after release. Mean migratory speeds between different receiver sites ranged from 0·49 to 1·82 body lengths (total length) per second (bl s⁻¹) for Atlantic salmon and 0·11–2·60 bl s⁻¹ for sea trout. Atlantic salmon were recorded 9, 48 and 77 km from the river mouth on average 28, 65 and 83 h after release, respectively. Sea trout were recorded 9 km from the release site 438 h after release. Only four (23%) sea trout were detected in the outer part of the fjord system, while the rest of the fish seemed to stay in the inner fjord system. The Atlantic salmon stayed for a longer time in the inner part than in the outer parts of the fjord system, but distinct from sea trout, migrated through the whole fjord system into the ocean.     

Daily feeding rhythms in Atlantic salmon I: feeding and aggression in parr under ambient environmental conditions

Daily feeding patterns of tank-reared Atlantic salmon Salmo salar parr under ambient conditions and their relationship with patterns of aggression were investigated during spring and autumn. A single, early afternoon feeding peak was observed during spring but no consistent pattern emerged from the autumn trial (fish did tend to feed in synchrony, even though the population's feeding pattern varied between days). Aggression was highest at first light during both spring and autumn, reducing thereafter throughout the day. Feeding and aggression were out of phase during spring, with the level of feeding increasing as aggression decreased. It is suggested that the high levels of early morning aggression are associated with re-establishment of the social order that breaks down after a period of darkness; in the spring this may have inhibited feeding.     

Seasonal and Diel Changes in Behaviour,Microhabitat use and Preferences by Young Pool-dwelling Atlantic Salmon,Salmo salar,and Brown Trout,Salmo trutta

There was a pronounced decline in activity of young pool-dwelling Atlantic salmon, Salmo salar, and brown trout, Salmo trutta, as the water temperatures dropped in the autumn and early winter, and the fish switched from a predominantly diurnal towards a nocturnal activity pattern. Such a switch in activity pattern has previously been observed in young brown trout, but the present study is the first documentation for juvenile Atlantic salmon under natural conditions. Juvenile fish fed actively even when water temperatures were below 0°C, although foraging behaviour at near-freezing temperatures was recorded exclusively during night surveys. This indicates that other proximate factors, in addition to water temperature, affect the activity of young salmon and trout in rivers. Trout kept feeding positions significantly higher above bottom than salmon in August and September, but both species reduced the height above bottom at the onset of winter, possibly due to reduced swimming performance and lowered food availability in the upper part of the water column.     

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.     

Evaluation of biochemical methods for the non‐destructive identification of sex in upstream migrating salmon and sea trout

Female‐specific markers of reproductive activity [plasma 17β‐oestradiol (E2), vitellogenin (VTG) and alkali‐labile phosphoprotein phosphorous (ALP)] were measured over 12 months in a captive population of brown trout Salmo trutta . During the early months of the reproductive season (February to May) and using the concentration of plasma E2 or plasma ALP as a marker for females the proportion of fish in which sex was misidentified was high (15–50%). The misidentification rate was considerably lower (1–8%) using plasma VTG. Preliminary evaluation of a commercial immunochromatographic VTG test system as a screen for the presence or absence of VTG in plasma from brown trout provided results that were consistent with those obtained from direct measurement of plasma VTG levels by enzyme‐linked immunosorbent assay (ELISA). These preliminary conclusions were verified by sampling upstream‐migrating anadromous brown trout, sea trout, and Atlantic salmon Salmo salar trapped over a 6 month period. Plasma E2 levels did not satisfactorily discriminate between male and female sea trout and Atlantic salmon. Plasma VTG levels in both species, however, were bimodally distributed and it was assumed that this divergence corresponded to male (plasma VTG levels <10 μg ml⁻¹) and female (plasma VTG levels >800 μg ml⁻¹) fishes. Plasma ALP provided a more accurate indication of sex in the wild Atlantic salmon and sea trout than was suggested by the pilot study on captive brown trout. The commercial immunochromatographic VTG test system provided results that were wholly consistent with the data obtained from the trapped fishes by direct measurement of plasma VTG.     

Effects of salmon lice infection and salmon lice protection on fjord migrating Atlantic salmon and brown trout post-smolts

Effects of artificial salmon lice infection and pharmaceutical salmon lice prophylaxis on survival and rate of progression of Atlantic salmon (n = 72) and brown trout post-smolts (n = 72) during their fjord migration, were studied by telemetry. The infected groups were artificially exposed to infective salmon lice larvae in the laboratory immediately before release in the inner part of the fjord to simulate a naturally high infection pressure. Groups of infected Atlantic salmon (n = 20) and brown trout (n = 12) were also retained in the hatchery to control the infection intensity and lice development during the study period. Neither salmon lice infection nor pharmaceutical prophylaxis had any effects on survival and rate of progression of fjord migrating Atlantic salmon post-smolts compared to control fish. Atlantic salmon spent on average only 151.2 h (maximum 207.3 h) in passing the 80 km fjord system and had, thus, entered the ocean when the more pathogenic pre-adult and adult lice stages developed. The brown trout, in comparison to Atlantic salmon, remained to a larger extent than Atlantic salmon in the inner part of the fjord system. No effect of salmon lice infection, or protection, was found in brown trout during the first weeks of their fjord migration. Brown trout will, to a larger extent than Atlantic salmon, stay in the fjord areas when salmon lice infections reach the more pathogenic pre-adult and adult stages. In contrast to Atlantic salmon, they will thereby possess the practical capability of returning to freshwater when encountering severe salmon lice attacks.     

Competitive habitat displacement of brown trout by Siberian sculpin: the role of size and density

Habitat competition in brown trout Salmo trutta and Siberian sculpin Cottus poecilopus was investigated by varying density, fish size, and species composition in stream channels providing areas of different substratum particle sizes. In allopatry, both small (52 ± 4 mm L _T) and large (86 ± 6 mm L _T) brown trout exhibited strong preference for the intermediate (8–11 cm diameter) and large (17–21 cm) gravel substrata. There was a tendency for more brown trout to occupy finer (2–4 cm) substrata with increasing density, in particular for large brown trout. Also, more small brown trout were observed on finer substrata when tested with large brown trout, suggesting interspecific competition for restricted space. Both small (56 ± 6 mm L _T) and large (88 ± 10 mm L _T) Siberian sculpin preferred the large gravel in all tests, and did not change their substratum preferences much with increasing densities, suggesting higher tolerance for 'crowding'. The large Siberian sculpin preferred the coarser substratum, and the largest individuals were consistently found on it. In sympatry with large Siberian sculpin, habitat displacement of brown trout occurred, indicative of interspecific competition. A higher proportion of small and large brown trout occupied the finer substrata than in allopatry. Habitat selection by large Siberian sculpin appeared to be unaffected by species composition and density. Small Siberian sculpin were displaced to finer substrata when tested with large Siberian sculpin, suggesting intraspecific competition. The results indicate that Siberian sculpin are potential habitat competitors for young brown trout.     

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.     

Production of juvenile salmonids in small Norwegian streams is affected by agricultural land use

1. We estimated the biomass and production of juvenile anadromous brown trout (Salmo trutta) and Atlantic salmon (Salmo salar) (parr) in 12 streams in the Skagerrak area of Norway to identify controlling environmental factors, such as land‐use and water chemistry. 2. Production estimates correlated positively with fish density in early summer, but not with the size of the catchment. The summer biomass of age‐0 brown trout and Atlantic salmon was smaller than that of age‐1 and constituted 27.4 and 25.7%, respectively, of the total biomass of the two groups. 3. Mean production of brown trout from July to September varied between streams, but in most cases it was below 2 g 100 m^?2 day^?1. Yearly cohort production from age‐0 in July to age‐1 in July was 10 g m^?2 or less, with mean annual production of 1.32 g 100 m^?2 day^?1, equivalent to 4.8 g m^?2 year^?1. The corresponding annual cohort production of Atlantic salmon was 0.38 g 100 m^?2 day^?1 or 1.4 g m^?2 year^?1. Annual production to biomass ratio (P/B) for brown trout of the same cohort in the various streams was between 1.47 and 4.37; the overall mean (±SD) for all streams was 2.25 ± 0.94. Mean turnover rate of Atlantic salmon was 2.73 ± 0.24. 4. Production of 0+ brown trout during the summer correlated significantly with the percentage of agricultural land and forest/bogs in the catchment, with maxima at 20 and 75%, respectively. Age‐0 brown trout production also correlated with concentration of nitrogen and calcium in the water, with maxima at 2.4 and 14 mg L^?1, respectively. 5. The results support the hypothesis that brown trout parr production reflects the quality of their habitat, as indicated by the dome‐shaped relationship between percentage of agricultural land and the concentration of nitrogen and calcium in the water.     

Clonal Atlantic salmon × brown trout hybrids produced by gynogenesis

Clonal full-sib progeny groups of Atlantic salmon Salmo salar × brown trout Salmo trutta hybrids were produced by gynogenesis. Eggs obtained from two 3-year-old Atlantic salmon (female) × brown trout (male) F₁ hybrids were activated with UV-irradiated rainbow trout Oncorhynchus mykiss sperm. Fecundity, percentage egg activation and percentage survival to completion of yolk-sac absorption were similar for the two females, and averaged 800 eggs kg⁻¹, 90 and 65%, respectively. Flow cytometric and protein electrophoretic analyses confirmed the progeny to be diploid hybrids. Isogenicity within progeny groups and to the maternal parent was indicated by identical DNA fingerprint patterns detected with multilocus oligonucleotide probes–GATA₍₅₎ and ACTG_(n). Isogenicity was also observed in the gynogenetic progeny of a third female spawned the following year. It appeared that a large portion of the oocytes in females of this hybrid underwent a premeiotic chromosome doubling, or possibly a complete suppression of meiosis. The result was ovulation of diploid eggs, each possessing a full set of both Atlantic salmon and brown trout chromosomes identical to those in the maternal somatic cells. Lines of clonal hybrids could therefore be perpetuated by gynogenesis and would have potential both as experimental animals and in commercial aquaculture.     

Within-stream variation in early life-history traits in brown trout

Significant additive genetic variance for most early life-history traits was found in brown trout Salmo trutta living in both allopatry above an impassable waterfall and sympatry (below the waterfall in the same stream) with alpine bullhead Cottus poecilopus. These traits included length, mass and yolk sac volume at hatching, and size at'button-up' (the time when yolk is enclosed within the body cavity). There were small differences in size at hatching and size at button-up among populations (adjusted for egg size). However, sympatric fry grew more rapidly and experienced lower mortality rates during the period of first feeding than allopatric fry. This might indicate behavioural differences between brown trout from the two populations. It is suggested that these phenotypic differences may be a result of adaptation to living in sympatry with alpine bullhead.     

Surface feeding and aggressive behaviour of diploid and triploid brown trout Salmo trutta during allopatric pair‐wise matchings

Diploid and triploid brown trout Salmo trutta were acclimated for 6 weeks on two feeding regimes (floating and sinking). Thereafter, aggression and surface feeding response were compared between pairs of all diploid, all triploid and diploid and triploid S. trutta in an experimental stream. In each pair‐wise matching, fish of similar size were placed in allopatry and rank was determined by the total number of aggressive interactions recorded. Dominant individuals initiated more aggression than subordinates, spent more time defending a territory and positioned themselves closer to the surface food source (Gammarus pulex), whereas subordinates occupied the peripheries. In cross ploidy trials, diploid S. trutta were more aggressive than triploid, and dominated their sibling when placed in pair‐wise matchings. Surface feeding, however, did not differ statistically between ploidy irrespective of feeding regime. Triploids adopted a sneak feeding strategy while diploids expended more time defending a territory. In addition, we also tested whether triploids exhibit a similar social dominance to diploids when placed in allopatry. Although aggression was lower in triploid pairs than in the diploid and triploid pairs, a dominance hierarchy was also observed between individuals of the same ploidy. Dominant triploid fish were more aggressive and consumed more feed items than subordinate individuals. Subordinate fish displayed a darker colour index than dominant fish suggesting increased stress levels. Dominant triploid fish, however, appeared to be more tolerant of subordinate individuals and did not display the same degree of invasive aggression as seen in the diploid and diploid or diploid and triploid matchings. These novel findings suggest that sterile triploid S. trutta feed similarly but are less aggressive than diploid trout. Future studies should determine the habitat choice of triploid S. trutta after release and the interaction between wild fish and triploids during the breeding season prior to utilization of triploids as an alternative management strategy within freshwater fisheries.     

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.     

Seasonality and heterogeneity of live fish movements in Scottish fish farms

Movement of live animals is a key contributor to disease spread. Farmed Atlantic salmon Salmo salar, rainbow trout Onchorynchus mykiss and brown/sea trout Salmo trutta are initially raised in freshwater (FW) farms; all the salmon and some of the trout are subsequently moved to seawater (SW) farms. Frequently, fish are moved between farms during their FW stage and sometimes during their SW stage. Seasonality and differences in contact patterns across production phases have been shown to influence the course of an epidemic in livestock; however, these parameters have not been included in previous network models studying disease transmission in salmonids. In Scotland, farmers are required to register fish movements onto and off their farms; these records were used in the present study to investigate seasonality and heterogeneity of movements for each production phase separately for farmed salmon, rainbow trout and brown/sea trout. Salmon FW-FW and FW-SW movements showed a higher degree of heterogeneity in number of contacts and different seasonal patterns compared with SW-SW movements. FW-FW movements peaked from May to July and FW-SW movements peaked from March to April and from October to November. Salmon SW-SW movements occurred more consistently over the year and showed fewer connections and number of repeated connections between farms. Therefore, the salmon SW-SW network might be treated as homogeneous regarding the number of connections between farms and without seasonality. However, seasonality and production phase should be included in simulation models concerning FW-FW and FW-SW movements specifically. The number of rainbow trout FW-FW and brown/sea trout FW-FW movements were different from random. However, movements from other production phases were too low to discern a seasonal pattern or differences in contact pattern.     

Marine food and diet overlap of co-occurring Arctic charr Salvelinus alpinus (L.), brown trout Salmo trutta L. and Atlantic salmon S. salar L. off Senja,N. Norway

Summary Stomach contents analyses and other biological information of Arctic charr (Savelinus alpinus (L.)), brown trout (Salmo trutta L.) and small Atlantic salmon (S. salar L.) caught 1982–85 close to the Åelv estuary (69°N) on the island of Senja, N. Norway are presented, and extracts of a 1975–85 fishing log given. this appears to be the first case study of the feeding habits of all three European anadromous salmonids in marine sympatry, and also one of very few reports on the marine food of the Arctic charr from Europe. The general feeding habits of the charr were similar to that found in N. Canada. Pelagic fish (herring, sand-eel) seem to be preferred. Plankton (crab megalopae, krill) and hyperbenthos (amphipods, mysids) are also taken, especially when suitable fish are scarce. In 1985 high herring densities provided superabundant food, and diet overlap between charr, trout and salmon was high. Salmonid nursery rivers are abundant in N. Norway and during summer the three species coexist in a near-shore, surface-oriented pelagic guild of fishes. The salmon seems to be a relatively specialized piscivore, while the trout takes a wider range of fish and also invertebrate prey. The charr probably is the most euryphagous of the three, being able to exploit the more marginal parts of the prey resources of their common habitat.