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     

Woody debris and terrestrial invertebrates – effects on prey resources for brown trout (Salmo trutta) in a boreal stream

Intensive forestry and other activities that alter riparian vegetation may disrupt the connectivity and the flux of energy between terrestrial and aquatic habitats and have large effects on biota, especially in small streams. We manipulated the amount of in-stream wood and the flux of terrestrial invertebrate subsidies to determine how these factors affected potential food resources for drift-feeding brown trout (Salmo trutta ) in a boreal Swedish forest stream. Specifically, we followed the effects on the abundance of aquatic and terrestrial invertebrate fauna from June to August 2007. The treatments were 1) addition of wood, unmanipulated terrestrial invertebrate inputs, 2) reduction of terrestrial invertebrate inputs (using canopy covers), no addition of wood, 3) unmanipulated ambient conditions, 4) simultaneous addition of wood and reduction of terrestrial invertebrate inputs. Added wood resulted in greater biomass of aquatic invertebrate biomass, and both input and drift of terrestrial invertebrates were reduced by canopy covers. In terms of total potential prey biomass, the addition of wood with ambient levels of terrestrial invertebrate inputs had the highest standing crop of benthic, wood-living and terrestrial invertebrates combined, whereas the treatment with reduced terrestrial input and no wood added had the lowest standing crop. Our study indicates that forest practices that both reduce the recruitment of wood and the input of terrestrial invertebrates to small streams have negative effects on prey availability for drift-feeding brown trout. The positive effects of wood addition on biomass of aquatic macroinvertebrates may partly compensate for the negative effects of reduced terrestrial invertebrate subsidies.     

Compensatory growth for free? A field experiment on brown trout,Salmo trutta

Laboratory studies suggest that animals may be capable of compensatory growth after periods of food shortage. There is, however, a lack of field experiments investigating the incidence and consequences of compensatory growth in the wild, and the relevance of compensatory responses in natural populations has recently been questioned. Here we addressed the hypotheses that (1) food restriction during critical growth periods can induce compensatory growth, and (2) that compensatory growth is associated with delayed costs in natural populations. These hypotheses were addressed by (1) manipulating the food intake of brown trout in spring, (2) measuring growth rate responses over the first month following release, and (3) measuring growth and mortality (i.e. recapture rate) over the subsequent fall and winter. We found that brown trout restored lost body weight and condition within a month, providing the first experimental demonstration of compensatory growth in the wild. However, no delayed costs of the compensatory response could be detected within the timespan of the experiment. We suggest that wild brown trout have an adapted "buffer capacity" to withstand fluctuations in food supply, allowing restoration of lost lipid reserves when feeding conditions improve. However, when prolonged food deprivation affect structural components, compensation may not be possible without compromising long-term performance.     

Does the trace element composition of brown trout Salmo trutta eggs remain unchanged in spawning redds?

The temporal stability of trace element concentrations in fertilized, artificially incubated anadromous brown trout Salmo trutta eggs and newly hatched fry was investigated. The anadromous status of the parental fish was confirmed using strontium isotopic analysis of otoliths. Whilst manganese concentrations in eggs varied over time, concentrations of aluminium, potassium, magnesium, strontium, barium and calcium were all unchanged 1 week and 6 weeks post‐fertilization as well as in recently hatched larvae. The results clearly suggest that the distinctive trace element signature present in the eggs and newly hatched larvae of anadromous S. trutta (typically characterized by high strontium, low barium) is stable over time. Therefore analysis of the trace element composition of eggs is concluded to be a cost‐effective and reliable method for determining the spatial and temporal extent of upstream spawning migration by anadromous salmonids. The temporal variability of at least one element in this study suggests the stability of untested multi‐element signatures cannot automatically be assumed.     

Utilization of brown trout Salmo trutta by Acanthocephalus clavula in an Irish lake: is this evidence of a host shift?

The population biology of the fish acanthocephalan Acanthocephalus clavula was described from 161 wild brown trout, Salmo trutta sampled over a two-year period in Clogher Lake in the west of Ireland. Overall prevalence of the parasite was 86% and the mean abundance was 53 worms per fish. Despite the presence of large numbers of worms in the trout very few females (2%) attained full reproductive maturity. This suggests that trout is an accidental host. A sample of yellow eels, Anguilla anguilla was examined at a different time from the same lake. The prevalence of A. clavula was 97% and the average abundance was 8 worms per fish. In contrast to the situation in trout, the proportion of female worms attaining full reproductive maturity was 61% fulfilling the expected characteristic of a preferred definitive host. The possible explanations for the very high abundance of A. clavula in trout are discussed and include the influence of fluctuations in host populations, host diet and the absence of a potential competitor.     

Seasonal and spatial variation in the diet of brown trout (Salmo trutta L.) in the subarctic River Laxá, north-east Iceland

This study examines seasonal and spatial patterns in the diet composition, stomach fullness and condition of a landlocked population of brown trout (2.5–61.3 cm) in the lake-fed River Laxá in NE-Iceland. The stomach contents consisted predominantly of benthic invertebrates, such as blackfly larvae Simulium vittatum (58%), chironomids (24%) and the freshwater snail Radix peregra (7%). The seasonal and spatial patterns of S. vittatum in the stomach contents were consistent with published studies on the life cycle and production of S. vittatum in the River Laxá. Close to the Lake Myvatn outlet (0–16 km), S. vittatum constituted 63% of the stomach content volume, and exhibited two peaks (i.e., generations) in its seasonal contribution to the stomach contents, whereas further downstream (16–35 km) S. vittatum constituted 45% of the volume and showed only clear evidence of one generation. Seasonal fluctuations in S. vittatum availability appear to affect the stomach fullness and the condition factor of the trout, especially close to the lake outlet where the fluctuations are pronounced.     

Is there genetic variation in the response to competition intensity in juvenile brown trout Salmo trutta?

Effects of intraspecific competition intensities on the relative performance (growth and movement) of juvenile brown trout Salmo trutta originating from nine different families were tested in tank experiments and in semi-natural streams. Both growth and movement differed consistently among families, indicating genetic variation in these traits. There were no significant interaction effects, however, between the intensity of competition and family origin on performance in either of the two experimental systems. Thus, genetic variation in response to competition intensity appeared to be limited in the population from which the juveniles used in this experiment originate.     

A 14-year study of habitat use and diet of brown trout (Salmo trutta) and Arctic charr (Salvelinus alpinus) in Lake Atnsjøen, a subalpine Norwegian lake

To reveal the mechanisms of sedimental H₂S accumulation, annual investigations on sedimental environments were conducted in two temperate estuarine lagoons. The lagoons, Gamo and Idoura (Japan), have similar shapes, locations, and topographical properties but different degrees of H₂S accumulation. Water stagnation causes a high phytoplankton biomass (Chl. a; 26–52 g l^–1) in the inner Gamo Lagoon. Gamo Lagoon sediment was characterized by high bounded sulfides (bounded Smainly FeS) and H₂S contents, and low C/N ratios (mean = 10.4) and iron (reactive Fe²⁺ and total Fe) contents. H₂S was not detected in Idoura Lagoon where phytoplankton biomass was much lower (Chl. a; 0.6–4 g l^–1). Idoura Lagoon sediment had high C/N ratios (mean=17.9) and high iron contents. The C/N ratio difference implies that organic matter in Gamo Lagoon originates mainly from more decomposable phytoplankton, while organic matter in Idoura Lagoon derives mainly from terrestrial vascular plants with lower decomposability. The excess loading of phytoplanktonic detritus in Gamo accelerates sedimentary microbial activity, including sulfate reduction (i.e., H₂S production). High Fe²⁺and low bounded S contents in Idoura sediment indicate a high chemical buffering capacity toward H₂S. In contrast, almost all Fe²⁺ in Gamo Lagoon had already reacted with H₂S as FeS. H₂S accumulation in Gamo Lagoon is caused by low sedimentary chemical buffering capacity toward H₂S, as well as higher microbial H₂S production, caused by the excess loading of phytoplanktonic detritus.     

Stocking impact and migration pattern in an anadromous brown trout (Salmo trutta) complex: where have all the stocked spawning sea trout gone?

We examined polymorphism at seven microsatellite loci among sea trout (Salmo trutta) (n = 846) collected from three areas in the Limfjord (Denmark). We then assessed their potential population source by comparing, using a mixed stock analysis (MSA) Bayesian framework, their genetic composition to that of brown trout collected from 32 tributaries pooled into nine geographical regions (n = 3801) and two hatcheries (n = 222) used for stocking. For each of the three regional sea trout groups (western, central and eastern Limfjord, n = 91, n = 426, n = 329, respectively), MSA was conducted with (i) all individuals in the group, (ii) with the subset of spawning sea trout only and (iii) with the subset of foraging, nonspawning individuals only, a subset that consisted primarily of sea trout caught during their first year at sea. For all three regional sea trout groups, a higher proportion of individuals (regardless of whether they were foraging or spawning) appear to have originated from the rivers that drain locally, than from the rivers that drain in other parts of the Limfjord. This suggests (1) that sea trout, at least during their first year at sea, undertake limited migrations within the Limfjord system and (2) that sea trout on their spawning run were caught close to their natal rivers. The proportion of sea trout of hatchery origin varied widely among all three Limfjord areas and broadly reflected regional stocking histories, with high proportions of sea trout of domestic origin in the east (39.3%), where stocking with domestic trout was practised intensely at the time of sampling, and in the west (57.2%), where a programme of coastal stocking of post smolts took place over several years in the early 1990s. In contrast, in the central Limfjord, where stocking with domestic trout was largely abandoned in the early 1990s, the proportion of sea trout of domestic origin was only 8.5%. Interestingly, for all three regional sea trout groups, virtually no sea trout of hatchery origin were found among the spawning individuals, which were on average larger than the nonspawning sea trout. These results suggest that stocked domestic brown trout that become anadromous experience high mortality at sea and are therefore largely absent among the larger, spawning individuals. We conclude that sea trout of domestic origin exhibit much reduced ability to reproduce and are unlikely to contribute significantly to the local gene pool largely because of a relatively high mortality at sea before the onset of maturity.     

Genetic variation at MHC, mitochondrial and microsatellite loci in isolated populations of Brown trout (Salmo trutta)

We have studied levels and distribution of genetic variation in nine isolated populations of Brown trout in NW Spain. In the present study, we have tried to test the importance of preservation of genetic variability for the survival of a set of isolated Brown trout (Salmo trutta) populations from the same river drainage. We screened genetic variation in three different markers, mitochondrial, microsatellites and Major Histocompatibility Complex (MHC), presumed to be under different selective pressures. Overall, genetic diversity varied considerably across populations and the distribution of genetic variation was similar at MHC and microsatellites; highly polymorphic populations at the microsatellite loci were also highly polymorphic at the MHC. We also observed high levels of differentiation among populations. Although we found evidence suggesting that balancing selection has influenced the long term evolution of the MHC, genetic drift seems to have eroded the effect of selection, becoming the predominant evolutionary force shaping genetic variation in some of the smaller populations. Despite current lack of variation at the MHC, these small populations seem to have remained viable for a long time.     

Salmo macrostigma (Teleostei,Salmonidae): Nothing more than a brown trout (S. trutta) lineage?

We examined specimens of the macrostigma trout Salmo macrostigma, which refers to big black spots on the flanks, to assess whether it is an example of taxonomic inflation within the brown trout Salmo trutta complex. Using new specimens, publicly available data and a mitogenomic protocol to amplify the control and cytochrome b regions of the mitochondrial genome from degraded museum samples, including one syntype specimen, the present study shows that the macrostigma trout is not a valid species. Our results suggest the occurrence of a distinct evolutionary lineage of S. trutta in North Africa and Sicily. The name of the North African lineage is proposed for this lineage, which was found to be sister to the Atlantic lineage of brown trout, S. trutta.     

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

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

Interspecific hybridization, a matter of pioneering? Insights from Atlantic salmon and brown trout

Interspecific hybridization may occur in situations of recent contact between a colonizer and a resident species, being more intense in the colonization front. Atlantic salmon Salmo salar and brown trout S. trutta have been sympatric species since their origin and they share spatial and temporal spawning niches, exhibiting low levels of bidirectional interspecific hybridization and introgression throughout their distribution range. Different causes have been identified for increased hybridization, from escapes or deliberate releases of domesticated fish to sneaking male behavior. We have examined hybridization rates and direction in different situations of advance of one of these species into a territory formerly inhabited by the other (247 samples were analyzed in northern Spain and 487 in Kerguelen Islands). In all cases, hybrids found in the colonization front were offspring of colonizer females and resident males. We hypothesize that these findings are the result of adaptive relaxed mate choice of colonizing females, regardless of the relative abundance of each species.     

Pex11α in brown trout (Salmo trutta f. fario): Expression dynamics during the reproductive cycle reveals sex-specific seasonal patterns

A negative correlation between female gonadal maturation kinetics and size variations of hepatic peroxisomes was earlier documented in brown trout, as a probable impact of serum estrogen changes during the reproductive cycle. Herein, we investigated whether the organelle volume/surface dynamics seen in female brown trout liver peroxisomes – without numerical changes within each hepatocyte – is followed by variations in the expression of the membrane peroxisome protein Pex11α gene. For comparison, we also studied males. We find in females a seasonal variation with the highest Pex11α expression in February, which was statistically different from all other tested periods. Overall, the expression of PEX11α had over a fivefold decrease from February to September. This period coincides with the reproductive transition between the earlier post-spawning gonadal remodeling and preparatory staging and the pre-spawning period. Males did not show changes. Our approach allowed the first characterization of a peroxin gene in a teleost, the Pex11α, while offering a correlation scenario were, as we hypothesized, the peroxisomal size kinetics is paralleled by membrane-related gene alterations (measured herein as proxy of Pex11α gene expression). Our data support and expand previous results on the regulation, function and morphology of peroxisome dynamics in brown trout, with a broader interest.     

Behavioural responses of female lake trout Salvelinus namaycush to male chemical stimuli and prostaglandin F2α

Male olfactory cues may guide aggregation on spawning reefs, mate evaluation and synchronized gamete release in lake trout Salvelinus namaycush, but a lack of information on the source and identity of the cues precludes direct tests of their function. Using a two-channel flume, we found ovulated female lake trout increased time spent in the channel treated with spermiated male-conditioned water, urine and bile but not synthesized prostaglandin F_2α. We suggest future efforts to characterize male olfactory cues focus on urine and postulate that bile acids contribute to its behavioural activity.     

Has stocking contributed to an increase in the rod catch of anadromous trout (Salmo trutta L.) in the Shetland Islands,UK?

The stocking of hatchery-origin fish into rivers and lakes has long been used in fisheries management to try to enhance catches, especially for trout and salmon species. Frequently, however, the long-term impacts of stocking programmes have not been evaluated. In this study, the authors investigate the contribution of a stocking programme undertaken to support the rod catch of sea trout in the Shetland Islands, UK. Once a highly productive recreational fishery, Shetland sea trout catches crashed in the mid-1990s. Around the time that stocking began, increases in rod catches were also reported, with advocates of the stocking highlighting the apparent success of the programme. Using a suite of genetic markers (microsatellites), this study explores the contribution of the stocking programme to the Shetland sea trout population. The authors found that the domesticated broodstock and wild spawned brown trout from seven streams were genetically distinct. Despite extensive stocking, wild spawned brown trout dominated, even in those streams with a long history of supplementation. The majority of sea trout caught and analysed were of wild origin – only a single individual was of pure stocked origin, with a small number of fish being of wild × stocked origins. This study suggests that stocking with a domesticated strain of brown trout has made only a very limited contribution to the Shetland Islands rod catch, and that the revival of sea trout numbers appears to be driven almost exclusively by recovery of trout spawned in the wild.     

The effects of short-term pH decrease on the reproductive output of the copepod Acartia bifilosa – a laboratory study

This laboratory study reports some reproductive responses of the copepod Acartia bifilosa to rapid variations in pH. The imposed changes mimic those that copepods could experience due to coastal upwelling, changed mixing conditions or vertical migration. We measured effects of low pH on egg production, hatching and early nauplii development (H₀: no effects on response variables between low and ambient pH). On treatment with low pH, we found positive effects on egg production rate and nauplii development time. The positive response to low pH could be an initial stress response or show that A. bifilosa is tolerant to the experimental pH values. The result suggests that A. bifilosa is adapted to pH changes as it performs daily migrations between the depths with differing pH. It could also be advantageous for population development if eggs hatch at high speed and so reduce the possibility that they will sink into anoxic and low pH waters.     

Haplotype (mtDNA) diversity of brown trout Salmo trutta in tributaries of the Austrian Danube: massive introgression of Atlantic basin fish — by man or nature?

Mitochondrial haplotype diversity in 27 populations of brown trout, Salmo trutta L., in Austria was investigated by sequencing the 5' end of the mitochondrial DNA (mtDNA) control region. Although all populations are within the Danube drainage, 44% of all individuals carried Atlantic basin haplotypes. It is argued that the presence of these haplotypes in Austria primarily reflects introgression stemming from the stocking of hatchery-reared fish. However, several lines of evidence suggest that some natural colonization from Atlantic lineages may have contributed to the present haplotype diversity. Nonetheless, the more diverse Danubian clade is represented by regionally distinct haplotype diversity that should be protected from the continued introduction of domesticated strains of exogenous fish     

Body Size, Diet and Growth of Landlocked Brown Trout, Salmo trutta, in the Subarctic River Laxá, North-East Iceland

Studies on diet preferences of stream-dwelling salmonids have mostly been limited to a relatively small range in body size. This study examined the influence of salmonid body size on prey size and diet composition, and its consequences for growth, in landlocked stream-resident brown trout, Salmo trutta (2.5–61.3cm) in the Laxá River, N-E Iceland. The most common prey of the 1622 trout sampled, were blackflies, Simulium vittatum, chironomid midges and the freshwater snail, Lymnaea peregra, which represented 56.3%, 21.8%, 10.8% of the stomach content volume, respectively. In general, the Laxá trout showed a consistent, but moderate, shift towards larger prey with increased body size. The relatively stable growth and the large body size attained were probably due to the high production of small benthic invertebrates and only secondarily to the ontogenetic shift towards larger prey.     

Is there a threshold size regulating seaward migration of brown trout and Atlantic salmon?

We investigated the influence of variation in body size and growth rate on age of smolting in Atlantic salmon and brown trout in four different Norwegian rivers. In Atlantic salmon smolt ages varied between 2 and 6 years, and in brown trout between 2 and 7 years. Smolt age was negatively correlated with parr growth, and positively correlated with smolt size. Age at smolting was more variable in the two northern than the two southern rivers. Smolt sizes and ages were also more variable in brown trout than in Atlantic salmon. Based on the observed variation in smolt size and age, we reject the hypothesis that a threshold size alone regulates age at smolting. Within populations smolt age depends on growth rate so that fast-growing parr smolted younger and smaller than slow-growing parr. We hypothesize that smolt size and age is a trade-off between expected benefits and costs imposed by differences in individual growth rate.