Stock‐specific variation of trophic position,diet and environmental stress markers in Atlantic salmon Salmo salar during feeding migrations in the Baltic Sea

This study investigated stock‐specific variation in selected ecophysiological variables during the feeding migrations of Atlantic salmon Salmo salar in the Baltic Sea. Oxidative stress biomarkers and EROD (ethoxyresorufin‐O‐deethylase, Cyp1A enzyme) activity were used as indicators of possible environmental stress and stable isotopes as determinants of diet and trophic position. Latvian S. salar stocks Daugava and Gauja had distinct stable‐isotope signatures compared to the other stocks, indicating differences in migration patterns, residency or arrival times, or dietary specialization among stocks. Salmo salar originating from Daugava and Gauja also had lower catalase enzyme activity than the other stocks. Post‐smolts originating from rivers of the Gulf of Finland had elevated EROD activities compared to fish of the same age from Bothnian Bay rivers, which could indicate exposure to organochlorine pollutants. No other stock‐specific differences in oxidative stress biomarkers were found. The study demonstrates how genetic, oxidative stress biomarker, EROD and stable‐isotope data may be combined to study trophic position, prey prevalence and environmental stress of mixed S. salar stocks foraging in the sea.     

Differentiation of western and eastern Baltic Sea cod stocks (Gadus morhua) by means of stable isotope ratios in muscles and otoliths

Stable isotope analysis of Baltic Sea cod from the Bornholm Sea (ICES‐SD 25) and western Baltic (Belt Sea, ICES‐SD 22) revealed significant differences in the δ¹⁵N and δ¹³C values of the dorsal muscle, as well as in the δ¹⁸O values of otoliths. The method pledges to become especially appropriate to the Baltic due to the high variability in oxygen isotope ratios associated with its estuarine nature.     

Comparison of the feeding apparatus and diet of European sardines Sardina pilchardus of Atlantic and Mediterranean waters: ecological implications

In this study, the feeding apparatus (gill rakers, GR) and the diet composition of European sardine Sardina pilchardus populations living in two contrasting environments were compared: the upwelling area off western Iberia and the comparatively less productive region of the north‐western Mediterranean Sea. The importance of local adaptations in the trophic ecology of this species was estimated. Sardina pilchardus from the Atlantic Iberian coast and from the north‐western Mediterranean Sea have clear differences in the feeding apparatus and diet compositions. Those from the Atlantic Iberian coast have significantly more GRs than S. pilchardus of the same size range in the Mediterranean Sea. While S. pilchardus from the Mediterranean Sea mostly depend on prey ranging between 750–1500 and 3000–4000 µm, corresponding mostly to cladocerans, decapods and copepods, those from the Atlantic depend on smaller prey (50–500 and 1000–1500 µm) that include phytoplankton and copepods, particularly during summer months, and S. pilchardus eggs during the winter. The marked difference between the trophic ecology of S. pilchardus in the two areas studied appears to have originated from different dietary strategies that the two populations have adopted in contrasting feeding environments. These differences are shown to profoundly affect the size and quality of prey consumed, and the effect of cannibalism on the populations.     

Evidence of Atlantic salmon Salmo salar fry movement between fresh water and a brackish environment

This study reports descent of Atlantic salmon Salmo salar fry from their natal streams to brackish waters of the Baltic Sea and their use of this environment as an alternative rearing habitat before ascending back to freshwater streams. To the authors' knowledge, residency in a brackish environment has not previously been demonstrated in S. salar fry. Recruitment success and evolutionary significance of this alternative life‐history strategy are presently not known.     

Environmental change influences the life history of salmon Salmo salar in the North Atlantic Ocean

Annual mean total length (L_T) of wild one‐sea‐winter (1SW) Atlantic salmon Salmo salar of the Norwegian River Imsa decreased from 63 to 54 cm with a corresponding decrease in condition factor (K) for cohorts migrating to sea from 1976 to 2010. The reduction in L_T is associated with a 40% decline in mean individual mass, from 2 to 1·2 kg. Hatchery fish reared from parental fish of the same population exhibited similar changes from 1981 onwards. The decrease in L_T correlated negatively with near‐surface temperatures in the eastern Norwegian Sea, thought to be the main feeding area of the present stock. Furthermore, S. salar exhibited significant variations in the proportion of cohorts attaining maturity after only one winter in the ocean. The proportion of S. salar spawning as 1SW fish was lower both in the 1970s and after 2000 than in the 1980s and 1990s associated with a gradual decline in post‐smolt growth and smaller amounts of reserve energy in the fish. In wild S. salar, there was a positive association between post‐smolt growth and the sea survival back to the River Imsa for spawning. In addition, among smolt year‐classes, there were significant positive correlations between wild and hatchery S. salar in L_T, K and age at maturity. The present changes may be caused by ecosystem changes following the collapse and rebuilding of the pelagic fish abundance in the North Atlantic Ocean, a gradual decrease in zooplankton abundance and climate change with increasing surface temperature in the Norwegian Sea. Thus, the observed variation in the life‐history traits of S. salar appears primarily associated with major changes in the pelagic food web in the ocean.     

Population and size-specific distribution of Atlantic salmon Salmo salar in the Baltic Sea over five decades

Population-specific assessment and management of anadromous fish at sea requires detailed information about the distribution at sea over ontogeny for each population. However, despite a long history of mixed-stock sea fisheries on Atlantic salmon, Salmo salar, migration studies showing that some salmon populations feed in different regions of the Baltic Sea and variation in dynamics occurs among populations feeding in the Baltic Sea, such information is often lacking. Also, current assessment of Baltic salmon assumes equal distribution at sea and therefore equal responses to changes in off-shore sea fisheries. Here, we test for differences in distribution at sea among and within ten Atlantic salmon Salmo salar populations originating from ten river-specific hatcheries along the Swedish Baltic Sea coast, using individual data from >125,000 tagged salmon, recaptured over five decades. We show strong population and size-specific differences in distribution at sea, varying between year classes and between individuals within year classes. This suggests that Atlantic salmon in the Baltic Sea experience great variation in environmental conditions and exploitation rates over ontogeny depending on origin and that current assessment assumptions about equal exploitation rates in the offshore fisheries and a shared environment at sea are not valid. Thus, our results provide additional arguments and necessary information for implementing population-specific management of salmon, also when targeting life stages at sea.     

Divergent trends in life‐history traits between Atlantic salmon Salmo salar of wild and hatchery origin in the Baltic Sea

Four Atlantic salmon Salmo salar stocks in the Baltic Sea, varying in their breeding history, were studied for changes in life‐history traits over the years 1972–1995. Total length (L_T) at age of captured (L_TC) fish had increased throughout the study period, partly due to increased temperature and increased L_T at release, (L_TR) but also due to remaining cohort effects that could represent unaccounted environmental or genetic change. Simultaneously, maturation probabilities controlled for water temperature, L_TC and L_TR had increased in all stocks. The least change was observed in the River Tornionjoki S. salar that was subject only to supportive stockings originating from wild parents. These results suggest a long‐term divergence between semi‐natural and broodstock‐based S. salar stocks. Increased L_T at age explained advanced maturation only marginally, and it remains an open question to what extent the generally increased probabilities to mature at early age reflected underlying genetic changes.     

Hybridization between Atlantic Salmon Salmo salar L. and Brown Trout S. trutta L. upon Artificial Propagation

Samples of Salmo salar and S. trutta were examined in 12 Russian fish hatcheries. With protein markers, hybrids of the two species were found in three hatcheries of the Baltic Sea basin. Some fishes had a phenotype intermediate between theS. salar and S. trutta phenotypes by morphological traits, but did not differ genetically from one of the parental species. Possible consequences of hybridization and ways to prevent it are discussed.     

Spawning-strategy-dependent diets in two North American populations of Atlantic salmon Salmo salar

The diet of repeat-spawner Atlantic salmon Salmo salar was investigated using carbon and nitrogen stable-isotope values from the outer growth band of scales, which reflect the fish's consumption and growth during their most recent marine phase. Isotope values for S. salar displaying different spawning strategies were compared between and within the Miramichi and Nashwaak Rivers, New Brunswick, Canada and a Bayesian mixing model was used to infer dietary contributions from potential prey items. Significant differences in the stable-isotope values were found among spawning strategies and between rivers, indicating differences in diet and feeding area, consistent with hypotheses. Bayesian mixing model results inferred the main prey items consumed during marine feeding by S. salar to consist of hyperiid amphipods and capelin Mallotus villosus for repeat alternate spawners from both rivers, sandlance Ammodytes sp. for repeat consecutive spawners from the Miramichi River and amphipods for repeat consecutive spawners from the Nashwaak River. These results demonstrate the diversity of feeding tactics among S. salar spawning strategies from the same river and between populations from different rivers. Accounting for differences in prey availability and the subsequent impact on S. salar diet and spawner return rates (i.e., marine survival) will facilitate the application of ecosystem-based management practices, such as ensuring that fisheries for forage species do not indirectly adversely affect S. salar return rates.     

Differences in fat accumulation between immature male and female Atlantic salmon Salmo salar after dietary administration of tetradecylthioacetic acid

This study provoked sex‐specific differences in fat metabolism in Atlantic salmon Salmo salar, by dietary administration of tetradecylthioacetic acid (TTA) during their first spring and winter in the sea. The effects of TTA were evaluated in June of the first spring and May of the second spring in the sea, by analysing white muscle‐fat content. Muscle fat in males and females differed significantly as a result of TTA in their diet and diet interacted with the sex of the fish. The fat content during the first spring after dietary TTA was lowered by a greater amount in females than in males, 3·1–4·3%, respectively (P < 0·05). In contrast, during the second spring, fat content was lowered by a greater amount in males than in females, 15·8–16·7%, respectively (P < 0·01). Condition factor followed a similar pattern to the muscle fat. The results indicate that the difference in male and female fat accumulation dynamics is related to sex‐specific reproduction biology of S. salar. In addition, the findings show that it is important to consider the sex of the fish and the season of the year when studying fat dynamics and reproductive biology of S. salar.     

The North Atlantic subpolar gyre and the marine migration of Atlantic salmon Salmo salar: the ‘Merry‐Go‐Round’ hypothesis

One model for marine migration of Atlantic salmon Salmo salar proposes that North American and southern European stocks (<62° N) move directly to feeding grounds off west Greenland, then overwinter in the Labrador Sea, whereas northern European stocks (>62° N) utilize the Norwegian Sea. An alternate model proposes that both North American and European stocks migrate in the North Atlantic Subpolar Gyre (NASpG) where S. salar enter the NASpG on their respective sides of the Atlantic, and travel counterclockwise within the NASpG until returning to natal rivers. A review of data accumulated during the last 50 years suggests a gyre model is most probable. Freshwater parr metamorphose into smolts which have morphological, physiological and behavioural adaptations of epipelagic, marine fishes. Former high‐seas fisheries were seasonally sequential and moved in the direction of NASpG currents, and catches were highest along the main axis of the NASpG. Marking and discrimination studies indicate mixed continental origin feeding aggregations on both sides of the Atlantic. Marked North American smolts were captured off Norway, the Faroe Islands, east and west Greenland, and adults tagged at the Faroes were recovered in Canadian rivers. Marked European smolts were recovered off Newfoundland and Labrador, west and east Greenland, and adults tagged in the Labrador Sea were captured in European rivers. High Caesium‐137 (¹³⁷Cs) levels in S. salar returning to a Quebec river suggested 62·3% had fed at or east of Iceland, whereas levels in 1 sea‐winter (SW) Atlantic Canada returnees indicated 24·7% had fed east of the Faroes. Lower levels of ¹³⁷ Cs in returning 1SW Irish fish suggest much of their growth occurred in the western Atlantic. These data suggest marine migration of S. salar follows a gyre model and is similar to other open‐ocean migrations of epipelagic fishes.     

Effects of hydropeaking on the spawning behaviour of Atlantic salmon Salmo salar and brown trout Salmo trutta

An in situ camera set‐up was used to study the spawning activity of Atlantic salmon Salmo salar and brown trout Salmo trutta throughout two consecutive seasons in a spawning area affected by hydropower‐related pulse flows due to hydropeaking. The purpose was to test whether the flow variation discouraged spawning in shallow areas or motivated spawning into areas with elevated risk of incubation mortality. There were more S. salar observed on the spawning ground during days with high discharge. The presence of S. salar in the spawning grounds was not affected by the hydropeaking cycles of the preceding night. Female S. salar were observed preparing nests within the first hour after water discharge had increased to levels suitable for spawning. In contrast, the number of S. trutta was not correlated with flow and nest preparation was also observed at a discharge corresponding to the lowest discharge levels during a hydropeaking cycle. Survival was generally high in nests excavated the following winter, with only 5·4% suffering mortality due to dewatering. The results suggest that S. salar may respond rapidly to variable‐flow conditions and utilize short windows with suitable flows for spawning. Smaller S. trutta may utilize low‐flow conditions to spawn in areas that are not habitable by larger S. salar during low flow.     

Variation in fat content and fatty‐acid composition of the Baltic herring Clupea harengus membras

The fat content and fatty‐acid profiles of herring, Clupea harengus membras, from the southern Baltic Sea varied depending on when (fishing season) and where (fishing grounds) the fish were caught as well as on their size and sex. The fat, protein and dry matter content and the fatty‐acid profiles were assayed in C. h. membras muscle tissue. The changes observed in fatty‐acid profiles were determined by factors such as specimen mass and fat content, which, in turn, depended on fishing season. This is explained by dietary differences between juvenile and older fish. Gonad maturation and spawning in the latter are also factors. The study results provide confirmation of the hypothesis that polyunsaturated fatty acids (PUFA), in particular docosahexaenoic acid (DHA) and eicosapentaenoic acid (EPA), play vital roles in the sexual maturation of C. h. membras.     

Fatty acid composition of sprat (<Emphasis Type="Italic">Sprattus sprattus</Emphasis>) and herring (<Emphasis Type="Italic">Clupea harengus</Emphasis>) in the Baltic Sea as potential prey for salmon (<Emphasis Type="Italic">Salmo salar</Emphasis>)

Sprat (Sprattus sprattus) and small herring (Clupea harengus) are the dominant prey fish of Atlantic salmon (Salmo salar) in the Baltic Sea. If the fatty acid (FA) proportions of sprat and herring differ, the dietary history of ascending salmon could be determined from their FA profiles. Therefore, we investigated the FA composition of several age groups of whole sprat and small herring, caught from the three main feeding areas of salmon in autumn and spring. Oleic acid (18:1n-9) was the most prevalent FA in sprat and characteristic of this species. In herring, palmitic acid (16:0) was the most common FA, but herring lipid was characterized by n-6 polyunsaturated FAs, and moreover, by palmitoleic acid (16:1n-7) and vaccenic acid (18:1n-7). Due to the higher lipid content of sprat, the concentrations of all other FAs, excluding these, were higher in sprat than in herring. The concentration of docosahexaenoic acid (DHA, 22:6n-3) increased with an increase in the lipid content and was consequently highest in the youngest specimens, being in young sprat almost double that of young herring, and 2.6 times higher in the sprat biomass than in that of herring. As a result of a decrease in the DHA concentration with age, the ratio thiamine/DHA increased with respect to age in both species, and was lower in sprat than in herring. It is concluded that an abundance of DHA in the diet of salmon most likely increases oxidative stress because of the susceptibility of DHA to peroxidation, and thus decreases thiamine resources of fasting, prespawning salmon. Because the FA composition of sprat and herring differs, and the relative abundancies of prey fish differ between the feeding areas of salmon, the feeding area of ascending salmon can most probably be derived by comparing their FA profiles.     

Stable carbon and nitrogen isotope ratios in muscle and epidermis of arctic whales

Collection of minimally invasive biopsy samples has become an important method to establish normal stable isotopes reference ranges in various wildlife species. Baseline data enhance the understanding of feeding ecology, habitat use, and potential food limitation in apparently healthy, free‐ranging cetaceans. Epidermis and muscle were collected from subsistence‐hunted northern Alaskan bowhead (n= 133 epidermis/134 muscle) and beluga whales (n= 42/49) and subsistence‐hunted Russian gray whales (n= 25/17). Additional samples were obtained from gray whales stranded in California (n= 18/11) during mortality events (1999, 2000). Both δ¹⁵N and δ¹³C are trophic position and benthic/pelagic feeding indicators, respectively, in muscle and epidermis. Epidermis is generally enriched in ¹⁵N over muscle, while epidermal ¹³C is more depleted. Lipid extraction does not alter δ¹⁵N in either tissue, but affects epidermal δ¹³C. Nitrogen‐15 is enriched in muscle, but not epidermis of stranded compared to subsistence‐hunted gray whales, indicating probable protein catabolism and nutritional stress in stranded whales. Similarly, epidermal δ¹³C of harvested whales is lower than in stranded whales, suggesting depleted lipid stores and/or food limitation in stranded animals. Epidermal isotope signatures are similar in both present‐day bowheads and in an ancient sample from the Northern Bering Sea region. Although only one specimen, this suggests trophic level of the ancient whale compares to modern bowheads after a millennium.     

Feeding habits of the whale shark (<Emphasis Type="Italic">Rhincodon typus</Emphasis>) inferred by fatty acid profiles in the northern Mexican Caribbean

Whale shark (Rhincodon typus, Smith, 1828) is an endangered species with anthropogenic pressures due to increasing demand of encounter tourism activities. Research efforts to identify management and conservation strategies for this species are needed. The Northern Mexican Caribbean is one of the most important feeding aggregation sites of whale sharks worldwide. In this study, Mexican Caribbean whale shark feeding habits are assessed by means of fatty acid (FA) signature analysis, a biochemical non-destructive technique widely applied in trophic ecology studies. Sub-dermal tissue biopsies of 68 whale sharks and samples of their potential prey (zooplankton) were collected during 2010 and 2011 in two areas with high R. typus abundance. Zooplankton samples (n?=?17) were divided in two categories: mixed zooplankton (several groups of zooplankton) and fish eggs (> 95% of sample components were fish eggs). FA profiles of whale shark tissue sampled between years showed significant variability; while there was no intraspecific differences in FA signature related to sex, size and location. FA profiles of whale sharks and their potential prey were dominated by saturated fatty acids (SFA). R. typus FA signature was significantly different from that of mixed zooplankton; on the other hand, whale shark and fish egg FA profiles formed groups with overlapping values and registered high levels of oleic acid. PUFA average ω3/ ω6 ratio on whale shark FA profiles for both years was below 1. Arachidonic acid (ARA) percentage was higher in whale shark biopsies (13.2% in 2010, 6.8% in 2011) compared to values observed in fish eggs (2.0%) and mixed zooplankton (1.4%). Similarity between FA profiles of whale sharks and fish eggs, low levels of bacterial FA found in R. typus biopsies, as well as whale shark feeding behavior observations in the study area, suggest that R. typus is feeding mainly on surface zooplankton in Mexican Caribbean; however, elevated ARA percentages in whale shark samples may indicate that this species has complementary feeding sources, such as demersal zooplankton, which has been reported in other aggregation sites. Results obtained contribute to the knowledge of the whale shark trophic ecology in the area, but are inconclusive. Further studies are recommended to evaluate whale shark FA profiles from different tissues (muscle or blood); also, broader information is needed about zooplankton FA signature in the study area.     

Phylogeographic evidence for the postglacial colonization of the North and Baltic Sea coasts from inland glacial refugia by Triglochin maritima L.

We investigated the geographical distribution of genetic variation in 67 individuals of Triglochin maritima from 38 localities across Europe using AFLP markers. Analysis of genetic variation resulted in the recognition of two major genetic groups. Apart from few geographical outliers, these are distributed (1) along the Atlantic coasts of Portugal, Spain and France and (2) in the North Sea area, the Baltic Sea area, at central European inland localities, the northern Adriatic Sea coast and the Mediterranean coast of southwest France. Considering possible range shifts of T. maritima in reaction to Quaternary climatic changes as deduced from the present-day northern temperature limit of the species, Quaternary changes of coastline in the North Sea area and the very recent origin of the Baltic Sea, we conclude that the coastal populations of T. maritima in the North Sea and Baltic Sea areas originated from inland populations.     

New insights into the trophic ecology of the scalloped hammerhead shark,Sphyrna lewini,in the eastern tropical Pacific Ocean

Scalloped hammerhead shark, Sphyrna lewini, is considered a generalist predator, showing individual specializations and different trophic levels. This species inhabits both oceanic islands and coastal nursery areas in the eastern tropical Pacific (ETP) Ocean. Existing trophic ecology studies on S. lewini have been focused on stomach contents. This study provides new insights into its foraging preferences, habitat use, and trophic position, using stable isotope analysis (SIA) of muscle tissue. Stable isotope signatures of δ¹³C and δ¹⁵N were determined in S. lewini muscles (n?=?29) from the Ecuadorian Pacific in 2013. Trophic position (TP), isotopic niches, and overlap of S. lewini were estimated by sex, age, and maturity stages to infer their dietary habits throughout life stages. SIA revealed complex movement patterns related to sex and age classes of S. lewini, highlighting high degrees of dietary plasticity and habitat use, with a stronger relation to coastal regions than previously reported. This study provides crucial information regarding essential areas and the related migration behavior of S. lewini, with important implications for their conservation and management in the ETP.

     

GENETIC STRUCTURE IN POPULATIONS OF FUCUS VESICULOSUS (PHAEOPHYCEAE) OVER SPATIAL SCALES FROM 10 M TO 800 KM1

Recent studies showing consequences of species’ genetic diversity on ecosystem performance raise the concern of how key ecosystem species are genetically structured. The bladder wrack Fucus vesiculosus L. is a dominant species of macroalga in the northern Atlantic, and it is particularly important as a habitat‐forming species in the Baltic Sea. We examined the genetic structure of populations of F. vesiculosus with a hierarchical approach from a within‐shore scale (10 m) to a between‐seas scale (Baltic Sea–Skagerrak, 800 km). Analysis of five microsatellite loci showed that population differentiation was generally strong (average F_ST = 12%), being significant at all spatial scales investigated (10¹, 10³, 10^4–5, 10⁶ m). Genetic differentiation between seas (Baltic Sea and Skagerrak) was substantial. Nevertheless, the effects of isolation by distance were stronger within seas than between seas. Notably, Baltic summer‐reproducing populations showed a strong within‐sea, between‐area (70 km) genetic structure, while Baltic autumn‐reproducing populations and Skagerrak summer‐reproducing populations revealed most genetic diversity between samples within areas (<1 km). Despite such differences in overall structure, Baltic populations of summer‐ and autumn‐reproducing morphs did not separate in a cluster analysis, indicating minor, if any, barriers to gene flow between them. Our results have important implications for management and conservation of F. vesiculosus, and we raise a number of concerns about how genetic variability should be preserved within this species.     

Influence of feed ration size on somatic and muscle growth in landlocked dwarf and farmed Atlantic salmon Salmo salar

We examined the possible adaptation of the dwarf Bleke population of Atlantic salmon Salmo salar from Lake Byglandsfjord in southern Norway to limited food resources. The growth performance and muscle development in juvenile Bleke and farmed S. salar under satiated or restricted (50%) feeding were examined for 10 months, starting 3 weeks after first-feeding stage. Four-thousand fish were divided into four replicated groups and random samples of 16–40 fish per group were measured six times during the experiment. The two strains showed no significant difference in mean body mass when fed restricted ration, but the individual variation was considerably higher in the farmed fish. Both Bleke and farmed S. salar grew significantly faster when fed to satiation, but the farmed S. salar showed much higher gain in mass and were three times heavier (201.5 g vs 66.7 g) and possessed twice as many fast muscle fibres (179,682 vs 84,779) compared with landlocked S. salar after 10 months. Farmed fish fed full ration displayed both hypertrophic and hyperplasic muscle growth, while the increased growth in Bleke S. salar was entirely associated with a larger fibre diameter. The landlocked Bleke strain has apparently adapted to low food availability by minimising the metabolic costs of maintenance and growth through reduced dominance hierarchies and by an increase in average muscle fibre diameter relative to the ancestral condition.