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

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

Current State of Conditions for Reproduction of the Black Sea Trout <Emphasis Type="Italic">Salmo trutta labrax</Emphasis> in the Mzymta River

Conditions of reproduction and distribution of the Black Sea trout Salmo trutta labrax in the basin of the Mzymta River after Olympic objects′ construction are considered. In the main river channel of the Mzymta, the structure of spawning and nursery grounds are destroyed along 57 km from the river mouth. The total area of these grounds in the Mzymta basin is 572800 m², and 49% of this area is situated above the dam of the Krasnopolyanskaya Power Plant and is inaccessible for spawners of the migratory form of the Black Sea trout listed in the Red Data Book of the Russian Federation. Consequently, natural reproduction of the migratory form discontinued. The resident form of trout still exists in the upper parts of the Mzymta and in 19 of its tributaries not affected by construction. On the 2.4-km-long stretch of the riverbed situated below the inflow of the Pslukh tributary, the substrate is in the process of natural restoration due to the descent of mudslides followed by washing of the substrate by the flow. The river is partially inhabited by trout individuals from habitats not affected by construction.     

Cytological status of the gonads and the level of thyroid and sex hormones in juvenile Black Sea trout,Salmo trutta labrax,of two phenotypic forms

Cytological state of gonads and hormonal status was studied in juvenile (15-month-old) Black Sea trout Salmo trutta labrax of the two phenotypic forms (resident and migratory). The differences in the rates of hametogenesis in the compared forms were revealed. It is shown that the spermatogenesis in the resident form is more intensive; on the other hand, the ovogenesis is slower. The levels of thyroxin and testosterone in blood are significantly higher in the groups of fish with intensively running gametogenesis. It is found that the concentration of triiodothyronine is slightly slower in the resident form than in the migratory. The correlation between levels of thyroid and sex steroid hormones is revealed.     

Experimental sea ranching of brook trout, Salvelinus fontinalis Mitchill

An experiment to induce anadromy in a population of wild brook trout, Salvelinus fontinalis , was conducted near Sept-Iles, Quebec, in 1978–1979. Brook trout were captured from the Matamek River, tagged and transported to the Matamek River estuary during late spring and early summer, and allowed free movement between an impassable waterfall 0.7 km upstream and the sea. Fish were recaptured in autumn as they returned to fresh water. Over two years, 34.0% of the released fish were recaptured. Best returns were in the 2+ and 3+ age classes with 38.0 and 62.1% recaptured, respectively. Straying of transplanted fish appeared to be <1%. All age classes included sea run brook trout (sea trout) but the largest percentages of sea trout occurred in older fish. Growth was better in sea trout than in fish which did not develop anadromy, presumably a function of an increased food supply at sea. Severe tagging effects stunted growth and probably suppressed anadromy, especially among younger fish. Sexual characteristics of recaptured fish indicated suppressed maturation of gonads in sea trout compared to fish remaining in fresh water and there was a shift to a larger percentage of females in the sea trout. Comparisons between our results and data on other anadromous Salvelinus species underscore the potential for sea-ranching of trout and char as a moderate effort, high yield aquaculture technique.     

Higher growth variability and stronger responses to temperature changes in wild than hatchery‐reared sea trout (Salmo trutta L.)

Each year, millions of hatchery‐reared sea‐run brown trout Salmo trutta L. (the sea trout) juveniles are released into the natural environment in the Atlantic region. The aim of this work was to investigate the growth responses of sea trout to changing temperature conditions and to compare the growth plasticity between wild and hatchery‐reared fish. Scales were collected from sea trout in a selected river flowing into the southern Baltic Sea. We analyzed the scale increment widths as a proxy of somatic growth and investigated the interannual variabilities and differences in growth between fish groups (wild and hatchery‐reared). We used mixed‐effects Bayesian modeling and ascribed the variances in growth to different sources. Furthermore, we developed indices of interannual (2003–2015) growth variation in the marine and freshwater phases of the life cycle of the fish and analyzed the relationships between trout growth and temperature. Temperature positively affects fish growth, regardless of the origin of the fish. We observed stronger relationships between fish growth and temperature conditions in the marine phase than in the freshwater phase. Additionally, wild sea trout are characterized by stronger responses to temperature variability and higher phenotypic plasticity of growth than those of the hatchery‐reared individuals. Therefore, wild sea trout might be better suited to changing environmental conditions than hatchery‐reared sea trout. This knowledge identifies possible threats in management actions for sea trout with an emphasis on ongoing climate change.     

Fatty Acid Status of Freshwater Resident and Anadromous Forms of Young Brown Trout (<Emphasis Type="Italic">Salmo trutta</Emphasis> L.)

No qualitative or quantitative differences in the profiles of fatty acids (FAs) belonging to the main classes have been found between freshwater resident (Orzega River, Onega Lake Basin) and anadromous (Indera River, White Sea Basin) forms of young brown trout. Differences have been registered in the conversion rate of the food-derived 18:2ω-6 and 18:3ω-3 FAs into long-chain and physiologically active FAs: 20:4ω-6 (arachidonic) and 22:6ω-3 (docosahexaenoic). Freshwater residential form of young brown trout (parr, at 3+-age) from the Orzega River are distinguished by a higher index of 20:4ω-6/18:2ω-6. Anadromous form of young of brown trout (smolts, at 4+-age) from the Indera River have a higher ratio of essential FAs 22:6ω-3/18:3ω-3 and ω-3/ω-6 polyunsaturated FAs; the last index correlates with higher weight–length characteristics of the studied individuals.     

Phylogeography and taxonomic status of trout and salmon from the Ponto‐Caspian drainages,with inferences on European Brown Trout evolution and taxonomy

Current taxonomy of western Eurasian trout leaves a number of questions open; it is not clear to what extent some species are distinct genetically and morphologically. The purpose of this paper was to explore phylogeography and species boundaries in freshwater and anadromous trout from the drainages of the Black and the Caspian Seas (Ponto‐Caspian). We studied morphology and mitochondrial phylogeny, combining samples from the western Caucasus within the potential range of five nominal species of trout that are thought to inhabit this region, and using the sequences available from GenBank. Our results suggest that the genetic diversity of trout in the Ponto‐Caspian region is best explained with the fragmentation of catchments. (1) All trout species from Ponto‐Caspian belong to the same mitochondrial clade, separated from the other trout since the Pleistocene; (2) the southeastern Black Sea area is the most likely place of diversification of this clade, which is closely related to the clades from Anatolia; (3) The species from the Black Sea and the Caspian Sea drainages are monophyletic; (4) except for the basal lineage of the Ponto‐Caspian clade, Salmo rizeensis, all the lineages produce anadromous forms; (5) genetic diversification within the Ponto‐Caspian clade is related to Pleistocene glacial waves; (6) the described morphological differences between the species are not fully diagnostic, and some earlier described differences depend on body size; the differences between freshwater and marine forms exceed those between the different lineages. We suggest a conservative taxonomic approach, using the names S. rizeensis and Salmo labrax for trout from the Black Sea basin and Salmo caspius and Salmo ciscaucasicus for the fish from the Caspian basin.     

Life history and genetic characterisation of sea trout Salmo trutta in the Adriatic Sea

1. The brown trout Salmo trutta is characterised by both anadromous (sea trout) and resident populations, naturally occurring in Atlantic and Ponto-Caspian rivers. Sea trout are currently considered absent from rivers of the Mediterranean area, probably because of the non-optimal chemical–physical characteristics of the Mediterranean Sea. However, the occasional bycatch of smoltified S. trutta in the Adriatic Sea is well known among fishermen and the biological explanation of this phenomenon is still controversial. The aim of this study was to compare the genetic diversity of freshwater and marine brown trout to try to understand the factors underlying the presence of putative anadromous brown trout in the Adriatic Sea.
2. In this study, we analysed the genetic diversity of: (1) wild brown trout collected from the Esino River (central Italy); (2) a domestic strain of brown trout used for stocking the study area; and (3) a sample of Adriatic sea trout collected near the outlet of the Esino River. Together with genetic analysis, we carried out scale analysis in order to track the freshwater/marine stages of the life cycle in the sea trout samples. The genetic characterisation was carried out by polymerase chain reaction–restriction fragment length polymorphism analysis of the mtDNA fragment ND-5/6 and the nuclear locus LDH-C1* and by genotyping 15 microsatellite loci. The genetic polymorphism obtained was used to investigate intra- and inter-population genetic diversity, rates of genetic introgression between wild and domestic samples and the origin of sea trout specimens by using assignment tests.
3. Our genetic analyses demonstrated that the sea trout analysed in this study are from the domestic strain of Atlantic origin used in central Italy for stocking activities. The level of genetic introgression between native and domestic samples is high in the Esino River. The populations more resilient to introgressive hybridisation appeared to be those living in the portion of the river network dominated by carbonate rocks. Assignment tests (GeneClass) suggest the existence of a link between stocking efforts and the freshwater origin of the sea trout. In addition, data obtained from the analysis of scales, size measurement, and sex determination showed a pattern of smolt age, size, and sex ratio very similar to those observed in other anadromous populations.
4. In conclusion, the present study highlighted that sea trout from the central Adriatic Sea originated from brown trout of Atlantic origin inhabiting the Esino River. Their seaward migratory behaviour could represent a consequence of an active migration instead of a passive displacement by water flow. Our results also showed that traditional stocking practices represent a negative activity for the conservation of the last Mediterranean native S. trutta populations.

     

Taxonomic status of salmonids in the Bulgarian stretch of the Danube River and their bionomic strategy

Analysis of salmonid fish samples from the Bulgarian stretch of the Danube resulted in the conclusion that this river segment is inhabited by the Black Sea salmon Salmo labrax Pallas, 1814 and not by the huchen Hucho hucho (Linnaeus, 1758), as originally believed. Occurrence of huchen in this stretch of the Danube River is improbable. It has also been suggested that the anadromous Black Sea salmon population does not exist. It remains to be determined if the salmon occurring in the Danube River and along the coast of the Turkish, Bulgarian, Romanian and Ukrainian coasts of the Black Sea represent the black trout, i.e. the brook form of the Black Sea salmon washed down from the tributaries or the specific potamodromous river form.     

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.     

Stocking experiments with 0 + and 1 + trout parr, Salmo trutta L., of wild and hatchery origin: 1. Post-stocking mortality and smolt yield

Wild and hatchery-reared 8–12-month-old (5–8 cm) trout, Salmo frurta L., were stocked in tributaries of the River Gudenb. Mortality was examined by means of electrofishing. Repeated electrofishing and handling caused a small increase in mortality. The daily instantaneous mortality rate Z was high during the first 2 months after stocking, ranging from 0.0070 for wild trout to 0.0326 for domestic trout at a stocking density of one trout per m² and from 0.0206 (wild trout) to 0.0888 (domestic trout) at a stocking density of two trout per m². Two months after stocking, Z decreased drastically ranging from 0.0007 (wild trout) to 0.0067 (domestic trout). When stocked, first-generation hatchery trout showed Z equal to domestic trout. Wild trout resident in the experimental stream were negatively affected by the introduction of domestic trout and wild trout from another stream. at a stocking density above the carrying capacity. It is concluded that the higher mortality of domestic trout was caused by changes in food, feeding and exercise, possibly combined with the lack of selection in the hatchery. Smolt yield at age 2+ was 3.2% (0+ trout stocked in the fall)-7.0% (1 + trout stocked in the spring) of the domestic trout stocked (approx. one-sixth to one-third of natural populations) and 65.2–68.7% of the domestic trout present before the smolt run. For first generation hatchery trout of wild origin the corresponding figures were 7.3% (age 0 +) and 93.4%, and for wild trout introduced to the experimental stream they were 11.1% (age0 +)and39.8%.     

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

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

A genetically distinct wild redband trout (<Emphasis Type="Italic">Oncorhynchus mykiss gairdneri</Emphasis>) population in Crane Prairie Reservoir,Oregon, persists despite extensive stocking of hatchery rainbow trout (<Emphasis Type="Italic">O. m. irideus</Emphasis>)

Crane Prairie Reservoir in the upper Deschutes River Basin has historically supported a wild population of migratory Deschutes River redband trout. Owing to its status as a premier destination for recreational angling in Oregon, the reservoir has been stocked with domesticated hatchery rainbow trout since 1955. In recent years the wild redband trout population has experienced a substantial decline. Effects on productivity related to genetic interaction with naturally spawning hatchery-origin fish (fitness risks) have not been determined. The species Oncorhynchus mykiss has been characterized with substantial genetic diversity throughout the Deschutes River Basin that further heightens the challenge of identifying specific conservation needs of wild populations. A conservation plan for Crane Prairie wild redband trout requires a better understanding of the natural reproductive success of out-of-basin hatchery trout in the reservoir tributaries, and the similarity between Crane Prairie redband trout with other extant redband trout populations in the basin. Using a suite of 17 microsatellite nuclear DNA markers, we evaluated the genetic structure among Crane Prairie Reservoir redband trout, hatchery rainbow trout, and two adjacent populations of redband trout from within the Upper Deschutes River Basin. We observed significant heterogeneity between the hatchery and wild Crane Prairie populations that may reflect differences in life histories, differential productivity and assortative mating. The genetic distinctions observed among the three redband trout populations suggest restricted gene flow and genetic drift within the upper basin. Temporally stratified sampling and larger numbers of samples will be necessary to confirm these conclusions.     

Distribution, growth and movement of River Usk brown trout (Salmo trutta)

The River Usk catchment in South Wales supports mainly freshwater resident brown trout, with few anadromous fish. Electric fishing and netting revealed that age-class distribution differed between main river and tributary habitats, the latter environment acting as a nursery area for young fish. Few fry were found at main river sites. Age-class distribution also differed between tributary systems, and possible reasons are discussed. Trapping experiments indicated that trout move to main river habitat at 2+ yr. Lengths at age (back-calculated from scale reading) were similar for main river and tributary resident fish at 1 and 2 year, but main river fish were larger at 3 and 4 yr. This habitat shift and size difference is discussed with reference to current angling regulations.     

Who are the missing parents? Grandparentage analysis identifies multiple sources of gene flow into a wild population

In order to increase the size of declining salmonid populations, supplementation programmes intentionally release fish raised in hatcheries into the wild. Because hatchery-born fish often have lower fitness than wild-born fish, estimating rates of gene flow from hatcheries into wild populations is essential for predicting the fitness cost to wild populations. Steelhead trout (Oncorhynchus mykiss) have both freshwater resident and anadromous (ocean-going) life history forms, known as rainbow trout and steelhead, respectively. Juvenile hatchery steelhead that 'residualize' (become residents rather than go to sea as intended) provide a previously unmeasured route for gene flow from hatchery into wild populations. We apply a combination of parentage and grandparentage methods to a three-generation pedigree of steelhead from the Hood River, Oregon, to identify the missing parents of anadromous fish. For fish with only one anadromous parent, 83% were identified as having a resident father while 17% were identified as having a resident mother. Additionally, we documented that resident hatchery males produced more offspring with wild anadromous females than with hatchery anadromous females. One explanation is the high fitness cost associated with matings between two hatchery fish. After accounting for all of the possible matings involving steelhead, we find that only 1% of steelhead genes come from residualized hatchery fish, while 20% of steelhead genes come from wild residents. A further 23% of anadromous steelhead genes come from matings between two resident parents. If these matings mirror the proportion of matings between residualized hatchery fish and anadromous partners, then closer to 40% of all steelhead genes come from wild trout each generation. These results suggest that wild resident fish contribute substantially to endangered steelhead 'populations' and highlight the need for conservation and management efforts to fully account for interconnected Oncorhynchus mykiss life histories.     

Net ground speed of downstream migrating radio-tagged Atlantic salmon (Salmo salar L.) and brown trout (Salmo trutta L.) smolts in relation to environmental factors

The downstream migration of Atlantic salmon (Salmo salarL.) and sea trout smolt (S. trutta L.) was investigated using radio telemetry in the spring of 1999 and 2000. Forty wild sea trout smolts, 20 F1 sea trout smolts, 20 hatchery salmon smolts and 20 salmon smolts from river stockings were radio tagged and released in the Danish River Lilleaa. The downstream migration of the different groups of fish was monitored by manual tracking and by three automatic listening stations. The downstream migration of radio tagged smolts of both species occurred concurrently with their untagged counterparts. The diel migration pattern of the radio tagged smolts was predominantly nocturnal in both species. Wild sea trout smolt migrated significantly faster than both the F1 trout and the introduced salmon. There was no correlation between net ground speed, gill Na⁺,K⁺-ATPase activity or fish length in any of the different groups. The migration speed of wild sea trout smolts was positively correlated with water discharge in both years. In F1 sea trout smolts, migration speed was positively correlated with temperature in 1999. The migration speed of salmon smolts did not correlate to any of the investigated parameters.     

The effect of stocking with 0+ year age‐class Atlantic salmon Salmo salar fry: a case study from the River Bush,Northern Ireland

An enhancement programme based on stocking 0+ year age‐class Atlantic salmon Salmo salar, conducted in the River Bush, Northern Ireland, U.K. over the period 1996–2005, was reviewed with reference to the performance and biological characteristics of wild fish. Wild ova to 0+ year fry (summer) survival was c. 8% with subsequent wild 0+ year fry‐to‐smolt survival c. 9%. Stocked unfed 0+ year juveniles gave c. 1% survival to smolt whilst fed 0+ year S. salar stocked in late summer exhibited survival at c. 5%. Stocking with unfed and fed fry contributed to increased smolt production and helped attain local management objectives between 2001 and 2005. Significant differences in biological characteristics were observed between wild and stocked‐origin fish. Wild‐smolt cohorts were dominated by 2+ year age‐class fish on the River Bush whilst smolts originating from fed fry mostly comprised younger 1+ year individuals. The mean mass of 1+ year smolts derived from stocked fed fry was significantly lower than that of wild 1+ year smolts, although these differences were not evident between older age classes. Differences in run timing between wild smolts and smolts derived from stocked fry were also apparent with the stocked‐origin fish tending to run earlier than wild fish. Although the stocking exercise was useful in terms of maximizing freshwater production, concerns over the quality of stocked‐origin recruits and the long term consequences for productivity are highlighted.     

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

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

Heart rate responses to predation risk in Salmo trutta are affected by the rearing environment

Both wild‐ and hatchery‐reared brown trout Salmo trutta , 18 months of age and of the same genetic origin, responded with increased heart rates (tachycardia) to a simulated predator attack on 2 consecutive days. Brown trout reared in the hatchery showed a more rapidly induced tachycardia compared with wild‐reared fish at day 1, but not day 2. During an undisturbed period several hours after attacks, hatchery‐reared brown trout maintained higher heart rates compared to wild‐reared fish on both days. Behavioural responses to the attack were very low for all fish, although hatchery‐reared fish tended to be more active than wild fish after the attack day 2. The observed differences may have had a genetic background caused by different selection regimes in the hatchery‐ and wild‐rearing environments, or could have been due to different phenotypic responses in the two environments.     

Rearing environment influences boldness and prey acquisition behavior,and brain and lens development of bull trout

Animals reared in barren captive environments exhibit different developmental trajectories and behaviors than wild counterparts. Hence, the captive phenotypes may influence the success of reintroduction and recovery programs for threatened and endangered species. We collected wild bull trout embryos from the Metolius River Basin, Oregon and reared them in differing environments to better understand how captivity affects the bull trout Salvelinus confluentus phenotype. We compared the boldness and prey acquisition behaviors and development of the brain and eye lens of bull trout reared in conventional barren and more structurally complex captive environments with that of wild fish. Wild fish and captive reared fish from complex habitats exhibited a greater level of boldness and prey acquisition ability, than fish reared in conventional captive environments. In addition, the eye lens of conventionally reared bull trout was larger than complex reared captive fish or same age wild fish. Interestingly, we detected wild fish had a smaller relative cerebellum than either captive reared treatment. Our results suggest that rearing fish in more complex captive environments can create a more wild-like phenotype than conventional rearing practices. A better understanding of the effects of captivity on the development and behavior of bull trout can inform rearing and reintroduction programs though prediction of the performance of released individuals.