Migration strategies of brown trout (Salmo trutta) in a subarctic river system as revealed by stable isotope analysis

We estimated the proportions of anadromous and freshwater‐resident brown trout (Salmo trutta) in different parts of the subarctic River Näätämöjoki/Neidenelva system (Finland and Norway) using carbon, nitrogen and hydrogen stable isotope analyses of archived scales as identifiers of migration strategy. Our results showed that carbon stable isotope values were the best predictor of migration strategy. Most individuals fell into two clearly distinct groups representing anadromous (47%) or freshwater‐resident (42%) individuals, but some fish had intermediate carbon values suggesting repeated movement between freshwater and the sea. The proportion of anadromous individuals decreased steadily with distance from the sea forming a spatial continuum in migration strategies which is probably maintained by the combination of several factors such as divergent availability of food resources, variable migration costs and genetic differences. These within‐catchment differences in migration strategies should be taken into account in fisheries management practices.     

Determining the spatial distribution of spawning by anadromous and resident brown trout Salmo trutta L using strontium content of eggs collected from redds

Abstract – Strontium (Sr) content of eggs collected from redds were used to determine the spawning contribution of resident and anadromous female brown trout in the Taieri River, New Zealand and were analysed on the catchment and tributary scale. At the catchment scale, analysis of Sr showed differences between eggs collected above and below the Taieri River Gorge. Samples collected above the gorge all had low‐Sr concentrations similar to levels found in an entirely resident brown trout population. Samples collected below the gorge exhibited a broad range in Sr concentrations, ranging from levels comparable to the known anadromous samples to levels comparable to the known freshwater‐resident samples. This suggests that the gorge prevents upstream migration by anadromous brown trout in the Taieri River. At the tributary scale, this pattern of anadromous brown trout spawning downstream of freshwater‐resident fish was repeated in one of the two tributaries located downstream of the gorge. Energetic cost of migration is the most likely explanation for the observed patterns in catchment and tributary scale distribution of spawning by resident and anadromous brown trout. It is concluded that the use of Sr concentrations in eggs collected from redds is a cost‐effective and reliable method to investigate the spawning contribution by anadromous brown trout.     

Environmental influences on life history strategies in partially anadromous brown trout (Salmo trutta,Salmonidae)

This paper reviews the life history of brown trout and factors influencing decisions to migrate. Decisions that maximize fitness appear dependent on size at age. In partly anadromous populations, individuals that attain maturity at the parr stage typically become freshwater resident. For individual fish, the life history is not genetically fixed and can be modified by the previous growth history and energetic state in early life. This phenotypic plasticity may be influenced by epigenetic modifications of the genome. Thus, factors influencing survival and growth determine life‐history decisions. These are intra‐ and interspecific competition, feeding and shelter opportunities in freshwater and salt water, temperature in alternative habitats and flow conditions in running water. Male trout exhibit alternative mating strategies and can spawn as a subordinate sneaker or a dominant competitor. Females do not exhibit alternative mating behaviour. The relationship between growth, size and reproductive success differs between sexes in that females exhibit a higher tendency to migrate than males. Southern populations are sensitive to global warming. In addition, fisheries, aquaculture with increased spreading of salmon lice, introduction of new species, weirs and river regulation, poor water quality and coastal developments all threaten trout populations. The paper summarizes life‐history data from six populations across Europe and ends by presenting new research questions and directions for future research.     

Reliance on lakes by salmon,trout and charr (Oncorhynchus,Salmo and Salvelinus): An evaluation of spawning habitats,rearing strategies and trophic polymorphisms

Salmonid fishes may reside within or migrate between stream and lake habitats, or undergo anadromous migrations between freshwater and the ocean. While the degree of anadromy of salmonids has been thoroughly compared, no analogous review has examined the degree of lake use. To assess the extent of reliance on lake habitat in this family, we considered 16 species of salmon, trout and charr from the genera Oncorhynchus, Salmo and Salvelinus, comparing their (a) use of lakes as spawning habitat, (b) rearing strategies in lakes, and (c) occurrence and diversity of lacustrine trophic polymorphism. In identifying the primary life‐history patterns of each species and exploring the lesser‐known lacustrine behaviours, we found that the extent of reliance on lakes exhibits a negative association with the degree of anadromy. Oncorhynchus rely least on lakes, Salmo to an intermediate level and Salvelinus the most, opposite of the general prevalence of anadromy among these genera. Lakes are critical to adfluvial and lake‐resident salmonids, but they also support anadromous and fluvial life histories by providing spawning, rearing, overwintering and/or summer refuge habitat. Adfluviality, although a non‐anadromous life history, consists of similar migration‐related traits and behaviours as anadromy, including the parr–smolt transformation, sex‐biased patterns of migration and residency, and the presence of precocious males. Lakes support life‐history variants, reproductive ecotypes and trophic morphs unique to lacustrine habitat. Therefore, conservation of salmonids is dependent on maintaining the diversity and quality of their habitats, including lakes.     

Population differentiation and temporal changes of carotenoid pigments and stable isotope ratios in the offspring of anadromous and non‐anadromous trout Salmo trutta

Trout (Salmo trutta) exhibit anadromous and non‐anadromous forms which are commonly sympatric. Offspring of the two forms can be separated by differences in characteristics such as stable isotope ratios and carotenoid pigments, which differ due to the influence of maternal resources. The rate of change in different characteristics due to freshwater feeding and the extent of differences between populations however remain unclear. Stable isotope (N and C) ratios and carotenoid pigment profiles were examined in the offspring of anadromous and non‐anadromous fish sampled at different times from six sites within the catchment of the River Tweed, UK. Both techniques were able to separate newly emerged fry successfully, with carbon isotopes distinguishing the forms better than nitrogen isotopes and zeaxanthin being the primary carotenoid pigment used to distinguish the offspring of different migratory forms. By 4 months, stable isotope ratios of the two forms were still distinct although both carbon and nitrogen ratios needed to be considered to distinguish the forms. Zeaxanthin levels were more variable and overlapped between the offspring of the two forms. There was significant variation between populations and over time in both isotope ratios and carotenoid composition. Comparison between the two techniques would suggest that stable isotopes are more effective for distinguishing between offspring of different forms as the distinctions are evident for longer. Population differences in isotope ratios could influence the extent to which the forms can be distinguished and need to be quantified more thoroughly to fully evaluate the technique.     

A survey of salmon gill poxvirus (SGPV) in wild salmonids in Norway

In 2016, the Norwegian health monitoring programme for wild salmonids conducted a real‐time PCR‐based screening for salmon gill poxvirus (SGPV) in anadromous Arctic char (Salvelinus alpinus L.), anadromous and non‐anadromous Atlantic salmon (Salmo salar L.) and trout (Salmo trutta L.). SGPV was widely distributed in wild Atlantic salmon returning from marine migration. In addition, characteristic gill lesions, including apoptosis, were detected in this species. A low amount of SGPV DNA, as indicated by high Ct‐values, was detected in anadromous trout, but only in fish cohabiting with SGPV‐positive salmon. SGPV was not detected in trout and salmon from non‐anadromous water courses, and thus seems to be primarily linked to the marine environment. This could indicate that trout are not a natural host for the virus. SGPV was not detected in Arctic char but, due to a low sample size, these results are inconclusive. The use of freshwater from anadromous water sources may constitute a risk of introducing SGPV to aquaculture facilities. Moreover, SGPV‐infected Atlantic salmon farms will hold considerable potential for virus propagation and spillback to wild populations. This interaction should therefore be further investigated.     

No difference in survival, growth and morphology between offspring of wild-born, hatchery and hybrid brown trout (Salmo trutta)

Abstract –  We studied survival, growth and morphological characters in the offspring of native hatchery and wild-born anadromous brown trout ( Salmo trutta ) and their hybrids (wild-born female × hatchery male and wild-born male × hatchery female) in a 1-year field experiment. We also conducted laboratory studies where we examined social interactions between the offspring of the same hatchery and wild-born trout. All offspring were raised in a hatchery and nose tagged before being released into the stream. In total, 1125 individuals were released into the stream (1999) and a total of 614 individuals were recovered (2000). We found no differences in growth and survival between the offspring of hatchery, wild-born and hybrid trout. Morphology was also similar among groups, where only 38% females and 36% males were classified into the right category, which were only 12% better than random classification. In the laboratory experiment, we compared only the offspring of hatchery and wild-born trout with respect to growth, dominance, aggressiveness, feeding and activity. We found small differences between the offspring of hatchery and wild-born fish with respect to growth but this effect was not found in the field experiment. Our result suggests that the offspring of hatchery trout and hybrids between hatchery and wild-born trout performed equally well to the offspring of wild-born trout.     

Do sea trout Salmo trutta parr surveys monitor the densities of anadromous or resident maternal origin parr,or both?

Brown trout Salmo trutta L. parr were sampled from 21 Estonian and three Finnish streams to investigate whether national sea trout parr surveys sample the progeny of anadromous or resident maternal parents. Otolith Sr:Ca core values were used and validated as a tool for distinguishing between the progeny of the two forms. In Estonia (n = 283), 92% of the parr were the progeny of anadromous maternal parents, and 8% were the progeny of resident maternal parents, whereas in Finland (n = 24), the respective proportions were 79% and 21%. Variation in the maximum otolith Sr:Ca core values among progeny of anadromous maternal parents indicated that some adult females may enter fresh waters several months before spawning. It was concluded that easily accessible locations situated up to 30 km from the sea largely contain progeny of sea trout, whereas sites with poor connectivity with the sea can be dominated by progeny of resident trout. This study demonstrated that the method applied provided an effective means to distinguish between the progeny of sea trout and resident brown trout.     

Genetic differentiation between anadromous and freshwater resident brown trout (Salmo trutta L.): insights obtained from stable isotope analysis

Abstract –  A genetic survey was carried out on recently emerging fry to investigate interbreeding between anadromous and freshwater resident brown trout in La Roche Brook, a major spawning tributary of the Oir River system (Normandy, France). Emerging fry were sampled at three different sites in spring 2002 and 2003. Stable isotope analysis (SIA) allowed the classification of each individual as the progeny of resident ( N  = 76) or anadromous ( N  = 58) female trout. Microsatellite DNA analysis showed that genetic diversity between fry samples can be explained by microgeographical (various sampling sites) and temporal (2 years of sampling) variations rather than maternal origin (anadromous vs. freshwater resident). We conclude that a high level of gene flow exists between the two morphs when anadromous adults have access to the spawning grounds of residents. We highlight the relevance of using SIA in conjunction with genetic analysis to study interactions between the two morphs.     

Growth and diet of anadromous Arctic charr after their return to freshwater

Abstract – The post smolt of anadromous Arctic charr ( Salvelinus alpinus ) from Lake Storvatn, northern Norway (70°39'48"), continued to feed and grow after their return to freshwater in late summer, but in late autumn (October) both their growth rate and feeding intensity ceased along with the drop in water temperature. In contrast, adult, sexually mature anadromous charr did not appear to feed during their freshwater residency. The stomach fullness of post smolts were similar to that of resident Arctic charr during early autumn, and their diets were similar, being dominated by pelagic prey such as surface insects, Holopedium sp., Bosmina sp. and chironomid pupae. Later in the autumn, the resident charr also included some more benthic prey, in particular gastropods, whereas the post smolts continued to feed predominantly on zooplankton and surface insects, but with a lower stomach fullness than the resident fish. This suggests that after arrival in freshwater, the post smolts continue their pelagic feeding behaviour displayed during the sea-migration.     

Winter movement activity patterns of anadromous Arctic charr in two Labrador lakes

Anadromous Arctic charr, Salvelinus alpinus, feed in the marine environment for several months during the summer and migrate back to fresh water in late summer to spawn and/or overwinter. While overwintering, anadromous Arctic charr are generally believed to reduce or cease feeding, and they are poorly described in their winter movement activity. This study used telemetry data collected from two locations to describe overwintering movement activity, including interindividual variation. Movement activity declined markedly during the ice‐covered period, suggesting opportunistic maintenance feeding was used as an energy conservation strategy. Fall and spring movement was correlated with daylight hours, and ice break‐up had a significant effect on the timing of outmigration. Movement activity was negatively correlated with body length, with smaller individuals being more active than larger fish. Although general activity patterns were evident, there were significant differences among individuals, particularly during spring immediately prior to lake departure. Lake size and individual differences in metabolic rate may account for some of this variation.     

Assigning origins in a potentially mixed‐stock recreational sea trout ( Salmo trutta) fishery

The anadromous, or sea‐going, life history form of brown trout, or sea trout ( Salmo trutta), may lead to potential mixing of populations while foraging at sea. In this article, we assess the potential that multiple populations are using common semi‐enclosed estuaries and quantify the potential levels of straying (i.e. dispersal) of foreign‐produced individuals into populations by using otolith chemical signatures as natural ‘tags’. To do so, we created a database of juvenile fish otolith chemistry (a marker of freshwater production) from four rivers and compared the chemistry of harvested fish in two estuaries important to anglers, the Renews River and Chance Cove Brook, to the database. A discriminant function analysis revealed significant differences in the otolith chemistry of juvenile fish inhabiting the four rivers with a 97% cross‐validated accuracy when classifying individual juveniles to their natal river, indicating our baseline was robust. When assigning adults caught over 3 years (2007–2009) in the recreational fishery in the Renews River estuary, it was determined that over 95% of the fish caught each year originated from Renews River. In contrast, harvested fish in Chance Cove during 2009 were disproportionately comprised of fish produced in Renews River, suggesting the potential for source‐sink population dynamics in Newfoundland. Taken as a whole, these results indicate limited population mixing in nearshore estuaries of this region, but also highlight the potential for some populations to subsidise the harvest by anglers in different areas.     

Parentally acquired differences in resource acquisition ability between brown trout from alternative life history parentage

Dominance hierarchies, where they exist, affect individual food acquisition ability and fitness, both of which have the potential to influence life history pathways. Juvenile salmonids exhibit clear dominance hierarchies in early life. As one of the drivers for the adoption of alternative life histories in salmonids is the relative rate of resource acquisition, there is potential for juvenile behaviour to influence the subsequent life history strategy of the individual. Lacustrine brown trout, Salmo trutta, exhibit a multitude of life histories which includes among others the piscivorous (ferox) life history where individuals grow to large size and have delayed maturity and benthivorous and pelagic life histories where individuals grow to much smaller sizes, however mature earlier. Using a number of observable characteristics of dominance, this study compared differences in behaviour between size‐matched pairs of progeny, reared under common garden conditions which are derived from alternative, co‐existing life history strategy parents. We found that first‐generation progeny of ferox trout were more aggressive, acquired more food, had lighter skin pigmentation and held more desirable positions than the progeny of benthivorous brown trout in an experimental stream system. Ferox trout progeny were dominant over benthivorous brown trout progeny in 90% of trials in dyadic contests. Given such clear differences in dominance, this study indicates that parentally acquired dominance‐related differences, passed through either, or both, of genetic and nongenetic (e.g. maternal effects) means, are likely a contributing factor to the continued maintenance of distinct life history strategies of brown trout.     

Summer survival and activity patterns of estuary feeding anadromous Salmo trutta

There is generally a very poor understanding of how anadromous brown trout, Salmo trutta, use marine ecosystems. In this study, we use acoustic telemetry techniques to test four alternative hypotheses for estuarine coastal habitat use by this species on a population in the Clyde, west-central Scotland. Anadromous brown trout in their second (or more) summer feeding in the marine environment did not show patterns of estuarine use predicted from studies on other populations of this species. They did not (a) use the inner Clyde Estuary as a staging post for onward migration to open coastal waters, nor did they (b) pass through the estuary rapidly nor did they (c) make multiple incursions into freshwater. In contrast, fish utilised a very geographically constrained area of the inner estuary (<8 km in length), within which fish were highly mobile, exhibiting more activity during a flooding tide and exhibited high survivorship compared with reports from other studies. The reasons why estuarine use by anadromous brown trout in the Clyde may differ from that reported elsewhere is discussed.     

Influence of maternal habitat choice,environment and spatial distribution of juveniles on their propensity for anadromy in a partially anadromous population of rainbow trout (Oncorhynchus mykiss)

This study evaluated the importance of the environment and spatial distribution of juvenile fish for the adoption of alternative migratory tactics in a partially anadromous population of rainbow trout (Oncorhynchus mykiss) from the Santa Cruz River. We captured young‐of‐the‐year fish along the river during autumn 2009, 2010 and spring 2010 and determined their maternal origin (anadromous vs. resident) using strontium to calcium ratios in the otolith core. Relative proportion of anadromous offspring, modelled with logistic regression, increased towards headwaters and in areas with deeper channels and larger substrate composition. Body length, modelled with linear multiple regression, varied positively with site depth, water velocity, substrate size and anadromous maternal origin. Based on evidence for limited juvenile movements (<25 km), the spatial extent of this study (240 Rkm) and the identification of large, contrasting reaches along the river, it is likely that the observed spatial distribution of juveniles and their association to sites with coarse substrate composition reflects maternal spawning activity. Results further indicate that anadromous females breed predominantly in middle and upper river sections in areas with coarse substrate. Given that body size in this system has been positively related to propensity for anadromy, we propose that female spawning choice affects their offspring's spatial distribution, providing the adequate physical template for anadromous offspring to reach or maintain larger body sizes and display anadromy themselves. Relevance of this study is also discussed in the context of alterations in response to future dam construction in one of the latest free‐flowing rivers sustaining anadromous O. mykiss.     

Water level influences migratory patterns of anadromous brown trout in small streams

Habitats modify the population ecology of species. Here, we show how low water level influences abundance and size of adult anadromous brown trout (Salmo trutta) entering a small, South Norwegian stream for spawning. After smolting, the fish appear chiefly to feed within 10 km of the home stream. In the autumn, South Norwegian streams typically flood because of heavy rainfall, when the anadromous brown trout entered from the sea. Mean annual duration of the upstream migration period was 34 days and ended when the flooding ended and the water temperature dropped to below 4°C. During most of the migration period, on average two trout ascended the river per day. The sexes entered the spawning area concurrently, and the male:female ratio of the anadromous trout was 1.27. No fish entered when the water depth just downstream of the spawning area was below 5 cm, and mean number of fish increased with increasing water depth to ca. 30 cm, but not at higher flows when the ascent gradually decreased. Mean and maximum size of the entering spawners increased with water depth between 5 and 16 cm. Among those that had been to sea, most were recaptured in the home stream, 4% in other streams, but only two of the strays were caught close to spawning time. The present results illustrate that population traits of anadromous brown trout from a small stream differ from those in larger rivers, probably because of selection associated with water flow.     

Anadromy and the dispersal of an invasive fish species (Oncorhynchus mykiss) in Eastern Quebec,as revealed by otolith microchemistry

Thibault I, Hedger RD, Dodson JJ, Shiao J‐C, Iizuka Y, Tzeng W‐N. Anadromy and the dispersal of an invasive fish species (Oncorhynchus mykiss) in Eastern Quebec, as revealed by otolith microchemistry. Ecology of Freshwater Fish 2010: 19: 348–360. © 2010 John Wiley & Sons A/S Abstract – Rainbow trout (Oncorhynchus mykiss) is invading rivers bordering the St. Lawrence Estuary (Quebec, Canada). Some rivers in Eastern Quebec support self‐sustaining populations while adult vagrants are frequently captured in rivers where no reproduction has been confirmed. We hypothesised that the development of anadromy has promoted the species dispersal. Otolith Sr:Ca analyses revealed that although all fish captured in the upstream stocking region were freshwater residents, both anadromous and freshwater resident phenotypes were found downstream in Eastern Quebec. The proportion of fish exhibiting the anadromous life cycle increased with the distance from the stocking zone. Eastern Quebec steelhead migrated to sea at the same age but at a larger size than steelhead within their native range. Age at first reproduction was similar to that observed in native populations. The development of the anadromous life cycle enables this species to colonise new rivers following long‐distance migrations along the St. Lawrence Estuary corridor.     

Novel insights into the marine phase and river fidelity of anadromous twaite shad Alosa fallax in the UK and Ireland

1. Most research on anadromous fishes has been invested in their freshwater life‐phases, resulting in a relatively sparse understanding of their spatial ecology during marine life‐phases. However, understanding the marine dispersal of anadromous fishes is essential to identify threats and to implement conservation measures that fully encompass their lifecycle.
2. The twaite shad Alosa fallax is an anadromous fish increasingly imperilled across its range due to pollution, harvesting, and impediments to freshwater migration, but little is known about its distribution and movements during its marine life‐phase. Here, the application of acoustic telemetry provided novel insights into the coastal dispersal of twaite shad in the UK and Ireland during 2018–2019, and the freshwater entry of individuals during the 2019 spawning season.
3. Of 73 twaite shad acoustic‐tagged during their upstream migration in the River Severn in May 2018, 58 emigrated from the river. Twelve were subsequently detected 200 km to the south‐west at the Taw–Torridge Estuary between July 2018 and April 2019, where estuarine movements up to 5.8 km inland occurred in summer, winter, and spring. One was subsequently detected in the Munster Blackwater Estuary (Ireland) and then in the River Severn, indicating a minimum movement distance of 950 km. Thirty‐four (59%) of the emigrating individuals from 2018 re‐entered fresh water in the rivers Severn (n = 33) and Wye (n = 2) in April and May 2019.
4. These results suggest year‐round use of estuarine and nearshore habitats by at least a subset of the twaite shad population during their marine phase, providing evidence of potential range overlap between populations that spawn in different areas in the UK and Ireland, which may be facilitated by substantial dispersal. The results also highlight the potential of telemetry for estimating freshwater and marine mortality, and the benefits of sharing detection data across networks.

     

Assessing the food web impacts of an anadromous Arctic charr introduction to a sub‐Arctic watershed using stable isotopes

Anadromous Arctic charr, Salvelinus alpinus (L.), was introduced to a sub‐Arctic river–lake system near the village of Kujjuuaq, Nunavik, and the stable isotope values and diets of key resident fish species were used to assess changes in feeding patterns. Stable isotope values for most species did not differ significantly between the pre‐ and post‐introduction periods, with observed shifts being within the bounds of expected natural variation. Lake chub, Couesius plumbeus (Agassiz), were the single species to show a difference between study periods, with a small but significant increase in δ¹⁵N. No significant post‐introduction changes were seen in lake trout, Salvelinus namaycush (Walbaum), omnivory or in any of the assessed quantitative food web metrics. Gut contents of major fish species similarly showed significant temporal overlap between the pre‐ and post‐introduction periods, and there was no significant change in species' weight–length relationships. The minor ecological impact was interpreted in relation to the availability of open niches exploitable by ecological generalists such as Arctic charr. The explanation accords with the known habitat and feeding flexibility of Arctic charr and the ecological immaturity of sub‐Arctic lakes known to have driven adaptive variation among Arctic charr. Findings suggest that anadromous Arctic charr may be introduced at moderate densities to other sub‐Arctic watersheds without major negative food web consequences for other resident fish species.     

An additional life-history tactic of masu salmon: Migration of parr to coastal habitats

Alternative life-history tactics of masu salmon Oncorhynchus masou are well documented. Subsequent to the freshwater parr stage (age ≥ 1 + years), an anadromous form migrates to the sea after smolting, while a resident form matures without seaward migration. In addition to this typical migratory dimorphism, anecdotal reports based on field observations have indicated that some underyearling masu salmon use estuarine waters. However, no empirical evidence indicates saltwater utilisation and subsequent survival in the early parr stage. Here, we used otolith microchemistry to examine whether a portion of masu salmon parr in northern Japan enters coastal habitats. The otolith Sr:Ca ratios of most juveniles collected from six rivers had consistently low values, indicating that masu salmon parr inhabiting these rivers stay only in freshwater. In contrast, in individuals from a steep-gradient river the Sr:Ca ratios increased at about a 200-µm distance from the otolith core. These results suggest that some masu salmon parr might use brackish water or sea water temporarily. In addition, three masu salmon parr were found in another steep river where a culvert located only ten metres from the river mouth completely blocked upstream migration for spawning. The Sr:Ca ratios in these fish increased at about >200 µm from the otolith core, indicating the parr had immigrated to the non-natal river from the sea. Such flexible behaviour at an early life stage may contribute to the spatial expansion of masu salmon, and the movement could moreover help to stabilise its population dynamics.