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.     

Distribution,prevalence and mean numbers of larvalAnisakis simplex (Nematoda: Ascaridoidea) in Atlantic salmon,Salmo salar L. and their use as biological indicators of host stocks

Synopsis An estimate of the arithmetic mean number of larvalAnisakis simplex per salmon,Salmo salar, in the north Atlantic was obtained from data collected off west Greenland where salmon from both sides of the Atlantic intermingle. An average of 5.34 larvae was found in the body cavity or associated viscera of 771 age 1 salmon representing some 60.8% of the total number of larvae present. LarvalA. simplex were found at all sampling stations throughout the north Atlantic. In most samples prevalence varied only slightly from area to area.The mean numbers of larvae in the body cavity and on associated organs of salmon in samples from North America were significantly different between samples. Comparisons of the mean numbers of larvae with several known variables in the data were, therefore, considered: host's sex, host's sea-age, geographic locality, year and season of capture.There was evidence that the mean number of larvae per host was greater in salmon from Europe than in those from North America, but the data could not be used to providereliable estimates of the mean numbers of larvae from each area because of heterogeneity within areas. Thus the data from west Greenland and from continental home waters could net be used in calculations to determine the percentage composition of North American and European salmon in the Greenland fishery although they supported earlier findings of mixing of salmon stocks off west Greenland.The use of parasitological data in the identification of different Atlantic salmon stocks is discussed.     

Movements of salmon Salmo salar (L.) to and from Irish waters

Fish tagged as kelts at a number of Irish stations have been recaptured outside Irish waters along the coasts of Great Britain, Norway and West Greenland. Salmon tagged as smolts in three Irish rivers have been recaptured off the coast of West Greenland and one off the Faroes. Sixteen fish tagged as smolts in Denmark, Great Britain and Sweden were recaptured in Irish rivers or on the coastline of Ireland. Clean salmon tagged in the open sea have travelled to the coasts of Great Britain and a single fish travelled to southern Sweden. Feeding salmon tagged off West Greenland, in the Labrador sea and off the Faroes have been recaptured in Irish waters.     

Development of seawater tolerance and subsequent downstream migration in wild and stocked young‐of‐the‐year derived Atlantic salmon Salmo salar smolts

This study investigated the development of hypo‐osmoregulatory capacity and timing of downstream migration in wild Atlantic salmon Salmo salar smolts from the River Stjørdalselva and stocked young‐of‐the‐year (YOY), derived S. salar smolts from the tributary River Dalåa. Both wild and stocked S. salar smolts developed seawater (SW) tolerance in early May, persisting through June, measured as their ability to regulate plasma osmolality and chloride following 24 h SW (salinity = 35) exposure. Although the majority of downstream migration among the stocked S. salar smolts occurred later than observed in their wild counterparts, the development of SW tolerance occurred concurrently. The wild S. salar from Stjørdalselva and stocked YOY smolts from the River Dalåa started to migrate on the same cumulative day‐degrees (D°). The study revealed no downstream migration before development of SW tolerance. This emphasizes the importance of incorporating physiological status when studying environmental triggers for downstream migration of S. salar smolts. Overall, these findings suggest that the onset of smolt migration in stocked S. salar smolts was within the smolt window from an osmoregulatory point of view.     

Evidence for episodic acidification effects on migrating Atlantic salmon Salmo salar smolts

Field studies were conducted to determine levels of gill aluminium as an index of acidification effects on migrating Atlantic salmon Salmo salar smolts in the north‐eastern U.S.A. along mainstem river migration corridors in several major river basins. Smolts emigrating from the Connecticut River, where most (but not all) tributaries were well buffered, had low or undetectable levels of gill aluminium and high gill Na⁺/K⁺‐ATPase (NKA) activity. In contrast, smolts emigrating from the upper Merrimack River basin where most tributaries are characterized by low pH and high inorganic aluminium had consistently elevated gill aluminium and lower gill NKA activity, which may explain the low adult return rates of S. salar stocked into the upper Merrimack catchment. In the Sheepscot, Narraguagus and Penobscot Rivers in Maine, river and year‐specific effects on gill aluminium were detected that appeared to be driven by underlying geology and high spring discharge. The results indicate that episodic acidification is affecting S. salar smolts in poorly buffered streams in New England and may help explain variation in S. salar survival and abundance among rivers and among years, with implications for the conservation and recovery of S. salar in the north‐eastern U.S.A. These results suggest that the physiological condition of outmigrating smolts may serve as a large‐scale sentinel of landscape‐level recovery of atmospheric pollution in this and other parts of the North Atlantic region.     

Retrospective growth analysis of Atlantic salmon Salmo salar and implications for abundance trends

Scale archives of Atlantic salmon Salmo salar from Maine, U.S.A., were examined to determine whether ocean conditions affected the long‐term trends in S. salar populations in the southern tier of the species' range in North America. To date, scale analyses of southern tier populations have been limited to hatchery fish; previous studies suggest that post‐smolt growth does not influence recruitment, with the exception that winter growth may play a role in stock maturation rate. A time series of scales from the Machias and Narraguagus Rivers spanning the years 1946 to 1999 was analysed. Image analysis was used to measure intercirculi spacing, which provided proxy variables of growth rate. Post‐smolt growth increment has increased since the early 1990s, as returns have decreased, suggesting that survival factors act on post‐smolts independent of growth. The data support the hypothesis of a decoupling between freshwater size and early marine growth. Growth during the second sea winter was independent of post‐smolt growth, suggesting that individuals are capable of significant compensatory growth. Southern tier North American stocks exhibit a similar pattern of independence between growth and survival as observed for northern tier North American stocks. These data support the inference that the recruitment of the North American and European subspecies is governed by fundamentally different mechanisms.     

Migration patterns and navigation cues of Atlantic salmon post-smolts migrating from 12 rivers through the coastal zones around the Irish Sea

The freshwater phase of the first seaward migration of juvenile Atlantic salmon (Salmo salar) is relatively well understood when compared with our understanding of the marine phase of their migration. In 2021, 1008 wild and 60 ranched Atlantic salmon smolts were tagged with acoustic transmitters in 12 rivers in England, Scotland, Northern Ireland and Ireland. Large marine receiver arrays were deployed in the Irish Sea at two locations: at the transition of the Irish Sea into the North Atlantic between Ireland and Scotland, and between southern Scotland and Northern Ireland, to examine the early phase of the marine migration of Atlantic salmon smolts. After leaving their natal rivers' post-smolt migration through the Irish Sea was rapid with minimum speeds ranging from 14.03 to 38.56 km.day⁻¹ for Atlantic salmon smolts that entered the Irish Sea directly from their natal river, to 9.69–39.94 km.day⁻¹ for Atlantic salmon smolts that entered the Irish Sea directly from their natal estuary. Population minimum migration success through the study area was strongly correlated with the distance of travel, populations further away from the point of entry to the open North Atlantic exhibited lower migration success. Post-smolts from different populations experienced different water temperatures on entering the North Atlantic. This was largely driven by the timing of their migration and may have significant consequences for feeding and ultimately survivorship. The influence of water currents on post-smolt movement was investigated using data from previously constructed numerical hydrodynamic models. Modeled water current data in the northern Irish Sea showed that post-smolts had a strong preference for migrating when the current direction was at around 283° (west-north-west) but did not migrate when exposed to strong currents in other directions. This is the most favorable direction for onward passage from the Irish Sea to the continental shelf edge current, a known accumulation point for migrating post-smolts. These results strongly indicate that post-smolts migrating through the coastal marine environment are: (1) not simply migrating by current following (2) engage in active directional swimming (3) have an intrinsic sense of their migration direction and (4) can use cues other than water current direction to orientate during this part of their migration.     

Passing a seawater challenge test is not indicative of hatchery‐reared Atlantic salmon Salmo salar smolts performing as well at sea as their naturally produced conspecifics

Despite satisfactory reactions to seawater challenge tests indicative of appropriate physiological state, hatchery‐reared Atlantic salmon Salmo salar smolts stocked in the Eira River in Norway between 2001 and 2011 performed less well at sea in terms of growth, age at maturity and survival than smolts of natural origin. The mean rates of return to the river for hatchery‐reared and naturally produced S. salar were 0·98 and 2·35%. In the Eira River, c. 50 000 hatchery‐reared S. salar smolts of local origin were stocked annually to compensate for reduced natural smolt production following regulation for hydroelectric purposes, while a mean of 17 262 smolts were produced naturally in the river. This study demonstrates that, although captive S. salar perform well in seawater challenge tests, hatchery‐reared smolts are not necessarily as adaptable to marine life as their naturally produced counterparts. These findings suggest that production of hatchery‐reared smolts more similar to naturally produced individuals in morphology, physiology and behaviour will be necessary to improve success of hatchery releases. Where possible, supplementary or alternative measures, including habitat restoration, could be implemented to ensure the long‐term viability of wild stocks.     

Post‐breeding movements of northeast Atlantic ivory gull Pagophila eburnea populations

The post‐breeding movements of three northeast Atlantic populations (north Greenland, Svalbard and Franz Josef Land) of the ivory gull Pagophila eburnea, a threatened high‐Arctic sea‐ice specialist, were studied between July and December 2007 using 31 satellite transmitters. After leaving their breeding grounds, all birds first dispersed eastward in August–September, to an area extending from the Fram Strait to the northwestern Laptev Sea (off Severnaya Zemlya). Most returned along the same flyway in October–November, hence describing a loop migration before moving south, off east Greenland. Wintering grounds were reached in December, in southeast Greenland and along the Labrador Sea ice‐edge, where Canadian birds also overwinter. One to two birds from each population however continued eastwards towards a third wintering area in the Bering Strait region, hence demonstrating a bi‐directional migration pattern for the populations and elucidating the origin of the birds found in the north Pacific during winter time. Overall, all birds breeding in the northeast Atlantic region used the same flyways, had similar rates of travel, and showed a peak in migratory activity in November. Though the total length of the main flyway, to the Labrador Sea, is only and at most 7500 km on a straight line, the mean total distance travelled by Greenland birds between July and December was 50 000 km when estimated from hourly rates of travel. Our study presents the first comprehensive and complete picture for the post‐breeding movements of the different ivory gull populations breeding in the northeast Atlantic.     

Evidence for long‐term change in length,mass and migration phenology of anadromous spawners in French Atlantic salmon Salmo salar

This study provides new data on Atlantic salmon Salmo salar life‐history traits across France. Using a long‐term recreational angling database (1987–2013) covering 34 rivers in three regions (genetic units), a decline in individual length, mass and a delayed adult return to French rivers was reported. Temporal similarities in trait variations between regions may be attributed to common change in environmental conditions at sea. The relative rate of change in phenotypic traits was more pronounced in early maturing fish [1 sea‐winter (1SW) fish] than in late maturing fish (2SW fish). Such contrasted response within populations highlights the need to account for the diversity in life histories when exploring mechanisms of phenotypic change in S. salar. Such detailed life‐history data on returning S. salar have not previously been reported from France. This study on French populations also contributes to reducing the gap in knowledge by providing further empirical evidence of a global pattern in S. salar across its distribution range. Results are consistent with the hypothesis that the observed changes in life‐history traits are primarily associated with environmental changes in the North Atlantic Ocean. They also emphasize the presence of less important, but still significant contrasts between region and life history.     

Basin‐scale phenology and effects of climate variability on global timing of initial seaward migration of Atlantic salmon (Salmo salar)

Migrations between different habitats are key events in the lives of many organisms. Such movements involve annually recurring travel over long distances usually triggered by seasonal changes in the environment. Often, the migration is associated with travel to or from reproduction areas to regions of growth. Young anadromous Atlantic salmon (Salmo salar) emigrate from freshwater nursery areas during spring and early summer to feed and grow in the North Atlantic Ocean. The transition from the freshwater (‘parr’) stage to the migratory stage where they descend streams and enter salt water (‘smolt’) is characterized by morphological, physiological and behavioural changes where the timing of this parr‐smolt transition is cued by photoperiod and water temperature. Environmental conditions in the freshwater habitat control the downstream migration and contribute to within‐ and among‐river variation in migratory timing. Moreover, the timing of the freshwater emigration has likely evolved to meet environmental conditions in the ocean as these affect growth and survival of the post‐smolts. Using generalized additive mixed‐effects modelling, we analysed spatio‐temporal variations in the dates of downstream smolt migration in 67 rivers throughout the North Atlantic during the last five decades and found that migrations were earlier in populations in the east than the west. After accounting for this spatial effect, the initiation of the downstream migration among rivers was positively associated with freshwater temperatures, up to about 10 °C and levelling off at higher values, and with sea‐surface temperatures. Earlier migration occurred when river discharge levels were low but increasing. On average, the initiation of the smolt seaward migration has occurred 2.5 days earlier per decade throughout the basin of the North Atlantic. This shift in phenology matches changes in air, river, and ocean temperatures, suggesting that Atlantic salmon emigration is responding to the current global climate changes.     

Population genetics ofAnisakis simplex (Nematoda: Ascaridoidea) in Atlantic salmon (Salmo salar) and their use as biological indicators of host stocks

Synopsis The extent to which data concerning the population genetics of a parasite — namely larvalAnisakis simplex, may be used as an auxilliary source of information on various aspects of the marine migrations of Atlantic salmon (Salmo salar) is examined.Frequencies of the various acid phosphatase alleles could not be used in distinguishing between or estimating the proportions of North American and European salmon occurring off west Greenland. The genetic structure of larvae from salmon taken in Scotland was different from that taken in Ireland and England confirming an earlier conclusion that the feeding habits of the fish from Scotland were significantly different. In the western Atlantic, salmon from northern Newfoundland and also salmon from the Bay of Fundy had apparently ingested different populations (or different proportions of the same nematode population) than salmon from Miramachi and Chaleur Bay areas. Samples from northern Newfoundland and Labrador were also genetically different. These findings are discussed in relation to the available literature concerning salmon migrations, explanations are put forward and areas of future research suggested.     

Phylogeography of North Atlantic intertidal tardigrades: refugia,cryptic speciation and the history of the Mid‐Atlantic Islands

Aim To analyse the phylogeographical history of intertidal tardigrades in the North Atlantic in order to improve our understanding of geographical differentiation in microscopic organisms, and to understand the potential importance of the Mid‐Atlantic Islands as stepping stones between the American and European coasts of the Atlantic Ocean. Location Twenty‐four localities from the Mid‐Atlantic Islands (Greenland, Iceland and the Faroe Islands) and both sides of the North Atlantic Ocean. Methods A mitochondrial marker (cytochrome c oxidase subunit I) was sequenced from individual tardigrades belonging to the genus Echiniscoides. The existence of cryptic species was detected using generalized mixed Yule coalescence analysis; lineage ages were estimated with relaxed clock methods; and the degree of geographical differentiation was analysed with samova analyses, haplotype networks and Mantel tests. Results Echiniscoides hoepneri, previously known only from Greenland, was recovered throughout the Mid‐Atlantic Islands. The Faroe Islands population was isolated from Greenland and Iceland, but overall genetic variation was low. The morphospecies Echiniscoides sigismundi had high genetic variation and consisted of at least two cryptic species. A northern and a southern species were both recovered on both sides of the Atlantic, but only the northern species was found on the Mid‐Atlantic Islands. The northern species showed signs of long‐term isolation between the Western and Eastern Atlantic, despite the potential of the Mid‐Atlantic islands to act as stepping‐stones. There was no sign of long‐term isolation in the southern species. The Mid‐Atlantic individuals of the northern species were of Eastern Atlantic origin, but Greenland and Iceland showed signs of long‐term isolation. The genetic pattern found in the southern species is not clearly geographical, and can probably be best explained by secondary contact between former isolated populations. Main conclusions North Atlantic intertidal tardigrades from the genus Echiniscoides showed strong geographical differentiation, and the Mid‐Atlantic Islands seemed unimportant as stepping stones across the Atlantic. The geographical variation of the northern species of E. sigismundi suggests post‐glacial recolonization from several refugia.     

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.     

Influence of sea temperature and initial marine feeding on survival of Atlantic salmon Salmo salar post-smolts from the Rivers Orkla and Hals, Norway

The abundance of returning adult Atlantic salmon Salmo salar, in the River Orkla in mid‐norway (1 sea‐winter, SW, fish) and River Hals in north Norway (1–3 SW fish), was tested against the early marine feeding and the seawater temperature experienced by their corresponding year classes of post‐smolts immediately after entry into the Trondheimsfjord (Orkla smolts, 22 years of data) and Altafjord (Hals smolts, 17 years of data). In both river–fjord systems, there was a significant positive correlation between the abundance of returning S. salar and the mean seawater temperature at the time of smolts descending to the sea. The number of 1SW fish reported caught in River Orkla was positively correlated to the proportion of fish larvae in the post‐smolt stomachs in Trondheimsfjord. The abundance of returning S.salar was, however, neither correlated to forage ratio (R_F) nor other prey groups in post‐smolt stomachs in the two fjord systems. In the Altafjord, the post‐smolts fed mainly on pelagic fish larva (70–98%) and had a stable R_F (0·009–0·023) over the 6 years analysed. In the Trondheimsfjord, however, there was a higher variation in R_F (0·003–0·036), and pelagic fish larvae were dominant prey in only two (50 and 91%) of the 8 years analysed. These 2 years also showed the highest return rates of S. salar in River Orkla. These results demonstrate that the thermal conditions experienced by post‐smolts during their early sea migration may be crucial for the subsequent return rate of adults after 1–3 years at sea. Pelagic marine fish larvae seem to be the preferred initial prey for S. salar post‐smolts. As the annual variation in abundance of fish larvae is related to seawater temperature, it is proposed that seawater temperature at sea entry and the subsequent abundance of returning adult S. salar may be indirectly linked through variation in annual availability of pelagic fish larvae or other suitable food items in the early post‐smolt phase.     

Chromosome polymorphisms track trans‐Atlantic divergence and secondary contact in Atlantic salmon

Pleistocene glaciations drove repeated range contractions and expansions shaping contemporary intraspecific diversity. Atlantic salmon (Salmo salar) in the western and eastern Atlantic diverged >600,000 years before present, with the two lineages isolated in different southern refugia during glacial maxima, driving trans‐Atlantic genomic and karyotypic divergence. Here, we investigate the genomic consequences of glacial isolation and trans‐Atlantic secondary contact using 108,870 single nucleotide polymorphisms genotyped in 80 North American and European populations. Throughout North America, we identified extensive interindividual variation and discrete linkage blocks within and between chromosomes with known trans‐Atlantic differences in rearrangements: Ssa01/Ssa23 translocation and Ssa08/Ssa29 fusion. Spatial genetic analyses suggest independence of rearrangements, with Ssa01/Ssa23 showing high European introgression (>50%) in northern populations indicative of post‐glacial trans‐Atlantic secondary contact, contrasting with low European ancestry genome‐wide (3%). Ssa08/Ssa29 showed greater intrapopulation diversity, suggesting a derived chromosome fusion polymorphism that evolved within North America. Evidence of potential selection on both genomic regions suggests that the adaptive role of rearrangements warrants further investigation in Atlantic salmon. Our study highlights how Pleistocene glaciations can influence large‐scale intraspecific variation in genomic architecture of northern species.     

Diversity and distribution of foliose Bangiales (Rhodophyta) in West Greenland: a link between the North Atlantic and North Pacific

Greenland is a continental island in the northern part of the North Atlantic where the foliose Bangiales flora is poorly known. It is an important area for the study of algal biogeography because of the region’s glacial history, in which Greenland has been alternately exposed to or isolated from the North Pacific via the Bering Strait. A molecular study using 3′ rbcL + 5′ rbcL–S sequences was undertaken to assess the diversity of foliose Bangiales on the west coast of Greenland and rbcL sequences were used to study the Greenland flora in a larger phylogenetic and floristic context. New and historic collections document seven species in four genera from the west coast of Greenland. All species had a close link to North Pacific species, being either conspecific with them or North Atlantic–North Pacific vicariant counterparts.     

Geolocator tagging reveals Pacific migration of Red‐necked Phalarope Phalaropus lobatus breeding in Scotland

The migration route of Red‐necked Phalarope populations breeding on North Atlantic islands has been subject to considerable speculation. Geolocator tags were fitted to nine Red‐necked Phalaropes breeding in northern Scotland to assess whether they migrated to Palaearctic or Nearctic wintering grounds. Of four birds known to return, two had retained their tags, of which one was recaptured. This male Phalarope left Shetland on 1 August 2012 and crossed the Atlantic Ocean to the Labrador Sea off eastern Canada in 6 days, then moved south to reach Florida during September, crossed the Gulf of Mexico into the Pacific Ocean and reached an area between the Galapagos Islands and the South American coast by mid‐October, where it remained until the end of April, returning by a similar route until the tag battery failed as the bird was crossing the Atlantic Ocean. The total migration of 22 000 km is approximately 60% longer than the previously assumed route to the western part of the Arabian Sea, and this first evidence of migration of a European breeding bird to the Pacific Ocean also helps to indicate the possible migratory route of the large autumn movements of Red‐necked Phalaropes down the east coast of North America.     

North Atlantic killer whale Orcinus orca populations: a review of current knowledge and threats to conservation

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