Migratory patterns of anadromous white-spotted charr Salvelinus leucomaenis in Eastern Hokkaido, Japan: The solution to a mystery?

We evaluated the migratory patterns of white-spotted charr in eastern Hokkaido, Japan using: (1) field observation of fish, (2) salinity-recording archival tags, and (3) analysis of otolith Sr and Ca concentrations. Field observations suggest that the majority of anadromous white-spotted charr outmigrate to the sea in spring (April to June) and ascend the river between late summer and autumn (August to November). Salinity records from archival tags revealed a fluctuating pattern consisting of both low and high values within a month, indicating frequent short-term movements between freshwater (including non-natal rivers) and brackish/salt water habitats. Otolith Sr:Ca ratios exhibited a sharp rise at about 40–50% of the otolith radius, which was followed by several spikes consisting of both low and high values, indicating that fish migrated between freshwater and marine environments multiple times during their life history. However, the peaks in Sr:Ca ratios were generally associated with annulus zones on the otolith, indicating that the anadromous migration was generally an annual event.     

Migratory timing, marine survival and growth of anadromous brown trout Salmo trutta in the River Imsa, Norway

The aim of the paper was to study sea migration, growth and survival of brown trout Salmo trutta of the River Imsa, 1976–2005. The migratory S. trutta were individually tagged and fish leaving or entering the river were monitored daily in traps located near the river mouth. The mean annual duration of the sea sojourn was 6–9 months for first-time migrants moving to sea between January and June. It was 8–18 months for those migrating to sea between July and December. Veteran migrants stayed 12 months or less at sea and most returned to the river in August. Early ascending fish stayed the longest in fresh water because most returned to sea in April to May. The day number of 50% cumulative smolt descent correlated negatively with mean water temperature in February to March and the February North Atlantic Oscillation index (NAOI). Mean annual sea growth during the first 2 years after smolting was higher for S. trutta spending the winter at sea than those wintering in the River Imsa. First year's sea growth was lower for S. trutta descending in spring than autumn. For first-time migrants, it correlated negatively with the February NAOI of the smolt year. Sea survival was higher for spring than autumn descending first-time migratory S. trutta with a maximum in May (14·9%). Number of anadromous S. trutta returning to the river increased linearly with the size of the cohort moving to sea, with no evidence of density-dependent sea mortality. Sea survival of S. trutta smolts moving to sea between January and June correlated positively both with the annual number of Atlantic Salmo salar smolts, the specific growth rate at sea, and time of seaward migration in spring. This is the first study indicating how environmental factors at the time of seaward migration influence the sea survival of S. trutta .     

Migrations of sea trout, Salmo trutta L., from the Vardnes river in northern Norway

During the periods 1956–1963 and 1967–1970 traps were operated to catch upstream- and downstream-migrating sea trout, Salmo trutta L. A total of 15 788 sea trout were tagged, using Carlin tags. The number of recaptures made in the traps was 4481, of which 1796 were recaptured more than once.
The distribution of the 2122 recaptures in the sea provides a picture of the sea-migration pattern. Of the sea recaptures, 52.8% were reported as within a distance of 3 km from the river mouth, compared to 0.7% more than 80 km away. All the different size-groups of sea trout were represented among both the long-distance and the short-distance migrants. The results of this study of sea trout migrations are discussed in relation to the published results for Atlantic salmon, Salmo salar L., and sea charr, Salvelinus alpinus (L.), from the same river.
The four highest values recorded for mean distance of daily travel away from the river were 20, 8, 8 and 6km day⁻¹ by smolts and 6, 6, 5 and 5km day⁻¹ by larger-sized sea trout.
Recaptures of tagged sea trout in rivers other than the Vardnes totalled 506, of which 306 had been tagged as smolts. The calculated minimum percentage of stray is 15.5%. The proportion of sea trout from the Vardnes river that actually spawn in other rivers is not known. No significant difference in length distribution was found between the sea trout caught in the Vardnes river and those caught in other rivers. An hypothesis concerning the selective advantages of straying by anadromous salmonids living in small rivers is discussed.     

Temperature requirements of Atlantic salmon Salmo salar, brown trout Salmo trutta and Arctic charr Salvelinus alpinus: predicting the effects of climate change

Atlantic salmon Salmo salar, brown trout Salmo trutta (including the anadromous form, sea trout) and Arctic charr Salvelinus alpinus (including anadromous fish) provide important commercial and sports fisheries in Western Europe. As water temperature increases as a result of climate change, quantitative information on the thermal requirements of these three species is essential so that potential problems can be anticipated by those responsible for the conservation and sustainable management of the fisheries and the maintenance of biodiversity in freshwater ecosystems. Part I compares the temperature limits for survival, feeding and growth. Salmo salar has the highest temperature tolerance, followed by S. trutta and finally S. alpinus. For all three species, the temperature tolerance for alevins is slightly lower than that for parr and smolts, and the eggs have the lowest tolerance; this being the most vulnerable life stage to any temperature increase, especially for eggs of S. alpinus in shallow water. There was little evidence to support local thermal adaptation, except in very cold rivers (mean annual temperature <6·5° C). Part II illustrates the importance of developing predictive models, using data from a long-term study (1967-2000) of a juvenile anadromous S. trutta population. Individual-based models predicted the emergence period for the fry. Mean values over 34 years revealed a large variation in the timing of emergence with c. 2 months between extreme values. The emergence time correlated significantly with the North Atlantic Oscillation Index, indicating that interannual variations in emergence were linked to more general changes in climate. Mean stream temperatures increased significantly in winter and spring at a rate of 0·37° C per decade, but not in summer and autumn, and led to an increase in the mean mass of pre-smolts. A growth model for S. trutta was validated by growth data from the long-term study and predicted growth under possible future conditions. Small increases (<2·5° C) in winter and spring would be beneficial for growth with 1 year-old smolts being more common. Water temperatures would have to increase by c. 4° C in winter and spring, and 3° C in summer and autumn before they had a marked negative effect on trout growth.     

Anadromous red-spotted masu salmon (Oncorhynchus masou ishikawae), a southernmost sea-migration form of salmonid,displays low variation in both age at seaward migration and sea age

We examined variations in age at seaward migration and sea age for the anadromous form of red-spotted masu salmon (Oncorhynchus masou ishikawae) in two Japanese rivers. The anadromous form of red-spotted masu salmon expressed only two sea migration patterns in the two rivers: (a) the majority of the salmon (95%, n = 81) were of age-0, and age-1 migrants were rare (n = 4); and (b) all the salmon examined (n = 22) made a return migration within a year, with 23% of the salmon exhibiting potamodromy in the river. Owing to low variation in their sea migratory patterns, the anadromous form of red-spotted masu salmon is likely vulnerable to environmental fluctuations.     

Habitat use and dispersal of post-smolt sea trout Salmo trutta in a Scottish sea loch system

Post-smolt anadromous brown trout Salmo trutta , sea trout, from two Scottish west coast rivers, the Balgy and Shieldaig, flowing into adjacent sea lochs were tracked simultaneously using arrays of moored acoustic receivers to determine dispersal patterns and loss rates. Fish tended to stay close to their natal rivers for the first 14 day after entering the sea, during which time about half the fish were lost to the study. Although initially the overall pattern of dispersal was similar for individual fish from both rivers, towards the end of the study the groups had converged into one of the loch basins. There were also pronounced individual differences in habitat use with all those fish detected for >42 days exhibiting different patterns of habitat use. Loss rates were similar between the two rivers despite differences in the range of air-breathing predators to which the fish were initially exposed. These findings suggest that any management of predators or other mortality agents should be targeted towards mouths of rivers during and immediately following smolt emigration.     

Pre-winter lipid stores in brown trout Salmo trutta along altitudinal and latitudinal gradients

Pre-winter lipid stores of brown trout Salmo trutta L. parr were compared along altitudinal (0-920 m a.s.l.) and latitudinal (58-71° N) gradients. There were increases in lipid content (size adjusted to common lipid-free dry mass of 2·0 g, corresponding to fresh mass of 10 g) with both increasing altitude and latitude. Mean size-adjusted lipid content for S. trutta in high altitude rivers was 60% higher than at low altitude (0·29 and 0·18 g, respectively). Mean size-adjusted lipid content for S. trutta in northern rivers was 30% higher compared to that in southern rivers (0·30 and 0·23 g, respectively). There was a marked between-river variation in mean lipid storage, probably reflecting different strategies or opportunities for the pre-winter acquisition of lipid both locally within rivers and between different populations. This study shows that temperature or winter length, not latitudinal covariates such as annual light regime, governs lipid storage patterns in juvenile salmonids.     

Dynamics of reproductive hormones during downstream migration in females of the Japanese eel, Anguilla japonica

The profiles of sex steroids (estradiol-17β, testosterone and 11-ketotestosterone) and the mRNA levels of gonadotropins (luteinizing hormone and follicle-stimulating hormone) were investigated before and after downstream migration in females of the Japanese eel species Anguilla japonica, which were collected in the brackish Hamana Lake and its inlet freshwater rivers. Eels were separated into three groups using otolith microchemistry: 'migrants' that grew in the inlet rivers and then made a downstream migration to Hamana Lake mainly in October and November; 'non-migrant' yellow eels caught in rivers during the same season; and 'residents,' which were yellow eels caught in rivers in August. Sex steroid levels, especially those of testosterone and 11-ketotestosterone, were higher in migrants than in non-migrants and residents. Real-time quantitative PCR analysis indicated that mRNA levels of luteinizing hormone (LH) β-subunits were significantly higher in migrants than in other groups, whereas those of follicle-stimulating hormone β-subunits did not show significant changes during downstream migration. The high levels of these hormones during downstream migration raise the question about if they also play a role in motivating the migratory behavior of eels.     

Archival tags reveal that Arctic charr Salvelinus alpinus and brown trout Salmo trutta can use estuarine and marine waters during winter

By tagging north-Norwegian anadromous riverine Arctic charr Salvelinus alpinus and brown trout Salmo trutta with simple archival tags that measured ambient water temperature and relating the recordings to the temperature in the surrounding environments, it was demonstrated that 91% of the S. trutta and 80% of the S. alpinus utilized the estuarine and marine environment during the winter. There was large individual variation in migratory behaviour. Salvelinus alpinus on average entered the estuary on 12 January and the marine environment on 26 February, had continuous marine migrations lasting up to 55 days, and spent on average 40 days in the estuary and 25 days in the sea during the winter. The corresponding numbers for S. trutta were 15 December and 4 January for first entry in the estuary and sea, maximum 39 days in the marine environment and average number of days in the estuary and sea were 34 and 50. Most individuals of both species frequently shifted between the three habitats. These findings thus contradict previous studies conducted on lake-dwelling populations, which suggested that northern populations of both species solely overwinter in fresh water. The use of inexpensive temperature recording archival tags gave detailed continuous information on seasonal migrations between habitats with different thermal characteristics.     

Life-history effects of migratory costs in anadromous brown trout

Mean size of sexually mature anadromous brown trout (sea trout) Salmo trutta in south-east Norway increased significantly with migratory distance ( D ) between the feeding area at sea and the spawning area in fresh water, from 32 cm for those spawning close to the river mouth to 43 cm at the spawning grounds 40 km inland. This was largely due to an increased size of the smallest anadromous spawners with increasing D . The raised mean size of the long-distance migrants is paralleled by an increased age at sexual maturity. Body mass at the same length of sea trout decreased with D in fresh water, meaning that the fish moving far inland was slimmer than those spawning near the coast. Gonadal mass of first-time spawning anadromous males declined significantly with D , and the fecundity and the ratio of fecundity over mean mass of the individual eggs adjusted for variation in fish mass, increased with D . There was no clear relationship between the ratio of anadromous to resident fish and D , probably because more variables than D , influence this relationship in the study streams.     

Identifying sea-run brown trout, Salmo trutta, using Sr : Ca ratios of otolith

 The migratory history of the brown trout, Salmo trutta, collected from Japanese rivers, was examined in terms of strontium (Sr) and calcium (Ca) uptake in the otolith, by means of wavelength dispersive X-ray spectrometry on an electron microprobe. Sea-run (anadromous) and freshwater-resident (nonanadromous) types of S. trutta were found to occur sympatrically. Otolith Sr concentration or Sr : Ca ratios of anadromous S. trutta fluctuated strongly along the life history transect in accordance with the migration (habitat) pattern from sea to freshwater. In contrast, the Sr concentration or the Sr : Ca ratios of nonanadromous fish remained at consistently low levels throughout the otolith. The higher ratios in anadromous S. trutta, in the otolith region from the core to 1500 μm, corresponded to the initial seagoing period, probably reflecting the ambient salinity or the seawater–freshwater gradient in Sr concentration. The findings clearly indicated that otolith Sr : Ca ratios reflected individual life histories, enabling the sea-run S. trutta to be distinguished from the freshwater-resident brown trout. Received: March 18, 2002 / Revised: May 15, 2002 / Accepted: June 5, 2002 Acknowledgments The authors are grateful to Messrs. T. Ikeda, S. Kudo, Y. Miyakoshi, M. Nagata, K. Shimoda, T. Takami, K. Takeuchi, and M. Ueda for their assistance in collecting specimens. This work was supported in part by Grant-in-Aid No. 13760138 from the Ministry of Education, Culture, Sports, Science and Technology, Japan. Correspondence to:Takaomi Arai     

Genomics and telemetry suggest a role for migration harshness in determining overwintering habitat choice,but not gene flow,in anadromous Arctic Char

Migration is a ubiquitous life history trait with profound evolutionary and ecological consequences. Recent developments in telemetry and genomics, when combined, can bring significant insights on the migratory ecology of nonmodel organisms in the wild. Here, we used this integrative approach to document dispersal, gene flow and potential for local adaptation in anadromous Arctic Char from six rivers in the Canadian Arctic. Acoustic telemetry data from 124 tracked individuals indicated asymmetric dispersal, with a large proportion of fish (72%) tagged in three different rivers migrating up the same short river in the fall. Population genomics data from 6,136 SNP markers revealed weak, albeit significant, population differentiation (average pairwise F_ST = 0.011) and asymmetric dispersal was also revealed by population assignments. Approximate Bayesian computation simulations suggested the presence of asymmetric gene flow, although in the opposite direction to that observed from the telemetry data, suggesting that dispersal does not necessarily lead to gene flow. These observations suggested that Arctic Char home to their natal river to spawn, but may overwinter in rivers with the shortest migratory route to minimize the costs of migration in nonbreeding years. Genome scans and genetic–environment associations identified 90 outlier markers putatively under selection, 23 of which were in or near a gene. Of these, at least four were involved in muscle and cardiac function, consistent with the hypothesis that migratory harshness could drive local adaptation. Our study illustrates the power of integrating genomics and telemetry to study migrations in nonmodel organisms in logistically challenging environments such as the Arctic.     

Migratory behaviour and growth of hatchery-reared post-smolt Atlantic salmon Salmo salar

Individually lagged, 1+ and 2+ hatchery-reared smolts of Atlantic salmon were released in spring and early summer at the mouth of the R. Imsa, south-western Norway. The post-smolts moved mainly northwards in the sea with the coastal current. The estimated mean migratory speed (± s.d. ) of those captured in the sea along the Norwegian coast was 7.45 (± 6.26) km day ⁻¹; in the fjords it was 1.63 (± 2.33) km day⁻². Many of the post-smohs ascended rivers the same year as released; 37.3% of the total number recaptured were caught in R. Imsa, upstream from the site of release, and 5.8% were caught in other rivers throughout middle and southern parts of Norway. The fish recaptured in rivers was probably sexually mature and entered rivers to spawn. Mean specific growth rate for post-smolts caught in the sea was higher than for those caught in R. Imsa (P <0.001) but not for those caught in other rivers (P> 0.05). Post-smolts ascending R. Imsa were smaller at release than those ascending other rivers. However, there was no size difference at release between post-smolts captured in the sea and those recaptured in rivers other than the R. Imsa.     

The biology, specificity and habitat of the species of Eubothrium (Cestoda: Pseudophyllidea), with reference to their use as biological tags: a review

The literature pertaining to the distribution, habitat, specificity, life cycles and population biology of all the species of Eubothrium is reviewed. Records in which scolex morphology is used to identify species cannot be used as this character is too variable to be of value in specific recognition. Study of the valid records suggests that E. crassum comprises three races. There is a freshwater race, whose preferred host is non-migratory Salmo trutta fario, but which also infects parr, smolts and landlocked adult S. salar and parr of migratory S. trutta trutta. Its distribution is confined to Europe and Eurasia and is co-extensive with that of S. trutta: it does not occur in N. America or E. Asia. It may be carried to sea in migratory salmonids, where it dies out in S. salar but may survive in S. trutta. Its life cycle only takes place in fresh water, and requires a copepod as the only intermediate host. Other fish species may serve as paratenic or accidental hosts. The preferred host of the marine Atlantic race of E. crassum is S. salar, but S. trutta is also infected. Its distribution is co-extensive with that of S. salar on both sides of the Atlantic Ocean. It may be carried into fresh water by both hosts, when its numbers decline in S. salar but not in S. trutta. The life cycle is unknown, but infection probably takes place in marine coastal waters. The preferred hosts of the marine Pacific race are all species of Oncorhynchus, with which its distribttion is co-extensive on both sides of the Pacific Ocean. It may also be carried into fresh water, and its life cycle is also unknown. E. salvelini comprises two races. The European freshwater race is specific to Salvelinus alpinus, and does not infect Salmo spp. Its life cycle takes place in fresh water and involves only one intermediate host, a copepod. It may be carried to sea in migratory fish, and survive to return to fresh water. The American race, found throughout North America and East Asia, infects species of Salmo, Salvelinus, Oncorhynchus, Cristivomer and other genera. Its biology is otherwise similar to that of the European race, although its return from the sea has not yet been confirmed in America. The other species of Eubothrium are more local in their distribution, some being endemic; do not comprise different races, and infect marine and anadromous fish. Nothing is known of their biology or life cycle. Eubothrium is recognised as being a marine genus that has invaded fresh water with anadromous fish. The inability to distinguish the races of E. crassum and E. salvelini on morphological characters, their low rate of survival in the other medium and their wide distribution renders them unsuitable as biological tags.     

Application of otolith microchemistry to estimate the migratory history of Japanese eel Anguilla japonica on the Sanriku Coast of Japan

The age and migratory history of the Japanese eel, Anguilla japonica Temminck & Schlegel, collected in Miyako Bay along the Sanriku coast of Japan, was examined using the otolith microstructure and analysis of strontium (Sr) and calcium (Ca) concentrations conducted with wavelength dispersive X‐ray spectrometry by an electron microprobe. The line analysis of Sr : Ca ratios along the life history transect of each otolith showed a peak (ca. 15–17 × 10^?3) which corresponded with the period of their leptocephalus and early glass eel stages in the ocean. The mean Sr : Ca ratios from the elver mark to the otolith edge indicated that there were eels with several general categories of migratory history, including sea eels that never entered freshwater (average Sr : Ca ratios, ≥6.0 × 10^?3), and others that entered freshwater for brief periods but returned to the estuary or bay. This evidence of the occurrence of sea eels in this northern area indicates that Japanese eels of the Sanriku coast do not necessarily migrate into freshwater rivers during recruitment as do glass eels at the beginning of their growth phase; even those that do enter freshwater may later return to the marine environment. Thus, anguillid eel migrations into freshwater are clearly not an obligatory migratory pathway, but rather a facultative catadromy with seawater or estuarine residents as an ecophenotype.     

Occurrence of sea eels of <Emphasis Type="Italic">Anguilla japonica</Emphasis> along the Sanriku Coast of Japan

 The age and migratory history of the Japanese eel, Anguilla japonica, collected along the Sanriku Coast of Japan, were examined using otolith microstructure and analysis of strontium (Sr) and calcium (Ca) concentrations. The mean Sr : Ca ratios from the elver mark to the otolith edge indicated that there were eels with several general categories of migratory history, including sea eels that never entered freshwater and others which had entered freshwater for brief periods but returned to the estuary or bay. This first evidence of the occurrence of sea eels in this northern area indicates that Japanese eels of the Sanriku Coast do not necessarily migrate into freshwater rivers. Received: May 15, 2002 / Revised: August 4, 2002 / Accepted: August 15, 2002 Acknowledgments We thank Messrs. S. Yamane and K. Morita, and crews of the Otsuchi Marine Research Center, Ocean Research Institute, The University of Tokyo, for their assistance in collecting the eels. This work was supported in part by Grant-in-Aid No. 13760138 from the Ministry of Education, Culture, Sports, Science and Technology, Japan. Correspondence to:Takaomi Arai     

Studies on the biology of Eubothrium salvelini and E. crassum in resident and migratory Salvelinus alpinus and Salmo trutta and in S. salar in North Norway and the islands of Spitsbergen and Jan Mayen

Samples of Eubothrium salvelini and E. crassum were obtained from resident and migratory Salvelinus alpinus and Salmo trutta and from S. salar at sea from North Norway and the islands of Spitsbergen and Jan Mayen. Scolex dimensions proved an unsuitable character for identifying these two parasite species, and use of alternative characters confirmed that E. salvelini is specific to Salvelinus and E. crassum to Salmo. Distributions of both species are co-extensive with their hosts, but levels of infection in any locality relate to the abundance of zooplankton. E. salvelini exhibits seasonal maturation and incidence cycles in char. It is recognised as being a freshwater parasite, although some individuals can survive the period of marine migration of sea char. Returning migrant char may contain such individuals together with small plerocerciform parasites acquired almost immediately upon their re-entry to freshwater. No evidence for marine infections with E. salvelini was found. The presence of plerocerciform and gravid adult E. crassum in salmon caught at sea confirms the existence of a marine life cycle and a marine race in this species. In salmon moving from one medium to the other the marine and freshwater races replace each other, but in sea trout, which spend shorter periods at sea, complete replacement may not occur and at any one time fish may harbour individuals of both races. Recognition of the existence of two races clarifies much of the confusion surrounding the biology of this species.     

Anadromy and marine habitat use of Lake trout (Salvelinus namaycush) from the central Canadian Arctic

Anadromy was documented in 16 lake trout, Salvelinus namaycush, from Canada's central Arctic using capture data and otolith microchemistry. For the first time, estuarine/marine habitat use was described for five individuals using acoustic telemetry. Age-at-first-migration to sea was variable (10–39 years) among individuals and most S. namaycush undertook multiple anadromous migrations within their lifetime. Telemetry data suggested that S. namaycush do not travel far into marine habitats and prefer surface waters (<2 m). These results further our collective understanding of the marine ecology of Arctic S. namaycush.     

Regional variation in otolith Sr:Ca ratios of African longfinned eel Anguilla mossambica and mottled eel Anguilla marmorata: a challenge to the classic tool for reconstructing migratory histories of fishes

Otolith Sr:Ca ratios of the African longfinned eel Anguilla mossambica and giant mottled eel Anguilla marmorata from nine freshwater sites in four rivers of South Africa were analysed to reconstruct their migratory life histories between freshwater and saltwater habitats. For A. mossambica, the Sr:Ca ratios in the otolith edge differed significantly among rivers and had large effect sizes, but did not differ among sites within a river. Otolith Sr:Ca ratios did not differ among rivers for A. marmorata. When rivers were pooled, the edge Sr:Ca ratios of A. mossambica were not significantly different from those of A. marmorata. According to the river-specific critical Sr:Ca ratio distinguishing freshwater from saltwater residence, most A. mossambica and A. marmorata had saltwater habitat experience after settlement in fresh water. This was primarily during their elver stage or early in the yellow eel stage. During the middle and late yellow eel stage, freshwater residency was preferred and only sporadic visits were made to saltwater habitats. The data also suggest that regional variations in otolith Sr:Ca ratios affect the critical Sr:Ca value and are a challenge for the reconstruction of migratory life histories that should be explicitly considered to avoid bias and uncertainty.     

Quantifying an overlooked aspect of partial migration using otolith microchemistry

For the first time, an overlooked aspect of partial migration was quantified using otolith microchemistry and brown trout, Salmo trutta, as a model species. Relative contributions of freshwater resident and anadromous female brown trout to mixed-stock sea trout populations in the Baltic Sea were estimated. Out of 236 confirmed wild sea trout sampled around the coast of Estonia 88% were of anadromous maternal origin and 12% were of resident maternal origin. This novel finding underscores the importance of the resident contingent in maintaining the persistence and resilience of the migratory contingent.