Predation on adult Atlantic salmon, Salmo salar L., by otters, Lutra lutra (L.), within the River Dee system, Aberdeenshire, Scotland

Predation on adult salmon, Salmo salur L., by otters, Lurra lutra (L.), varied seasonally on the R. Dee, Aberdeenshire, Scotland, being highest during the spawning season in winter. Predation is described for some tributaries of the river. Male fish were caught by otters more often than females, and it is suggested that they were most vulnerable during their extensive excursions up and down stream, particularly as they negotiated shallow riffles. Otters appeared to prey upon healthy fish rather than those infested with Saprolegnia sp. but there was no difference in the size, freshwater- or sea-age offish killed by otterscompared with 'kelts' which had died non-violently. At least some of the otters obtained most of their daily food requirements by catching a single salmon per night.
Considerable numbers of adult salmon may be killed by otters during the spawning period but it is suggested that, because most are males, this is unlikely to affect the breeding success of the salmon population. Most predation occurs outside the fishing season and so is unlikely to reduce the numbers of salmon caught by anglers.     

Salmon lice, Lepeophtheirus salmonis, infestation as a causal agent of premature return to rivers and estuaries by sea trout, Salmo trutta, juveniles

A field experiment conducted in the River Lønningdalselven in spring 1992 supports the hypothesis that salmon lice, Lepeophtheirus salmonis, infestations may cause premature return of sea trout, Salmo trutta, juveniles, either to estuaries or to rivers. When lice infested (exposed) and uninfested (control) sea trout juveniles (post smolts) were released simultaneously into the sea, exposed fish returned to the estuarine area earlier compared with controls. Within the following two days, exposed sea trout migrated further into freshwater. At that time they were infested with a median of 62.5 lice, dominated by chalimus larvae and late juveniles. Exposed sea trout suffered from an osmoregulatory failure in sea water and this is considered one reason for infested fish returning to brackish water. While only a few control fish returned to the estuary on the day of release, some more returned to freshwater the following four days. During this time they had become heavily infested with copepodids, and carried a median of 150.0 lice. It is suggested that physiological stress and high infection pressure in the sea results in sea trout juveniles returning to estuaries and freshwater.     

Upstream migration of adult Atlantic salmon past a fish counter weir in the Aberdeenshire Dee, Scotland

During their upstream migration through the lower reaches of the Aberdeenshire Dee, Scotland, radio-tagged adult Atlantic salmon passed a Crump weir-based fish counter without substantial delay over a range of river flows and water temperatures. Tagged salmon were detected sequentially by automatic listening stations placed along the river and were detected at the counter site for a median time of 18 min, compared with a median time of 5 min before the weir was built. One fish out of 16 spent 6·7 h in the vicinity of the weir, but such migratory pauses also occurred before the weir existed. The results suggest that the number of salmon crossing the counter per unit time is a function of the number of fish present downstream and their tendency to migrate upstream, without being limited by their ability to surmount the weir. The apparent slight delay at the weir is probably insignificant in terms of the overall progress of riverine migration and in the context of using counter records to relate salmon movements to environmental Variables.     

Does increased abundance of sea lice influence survival of wild Atlantic salmon post‐smolt?

A synthesis of results from two projects was assessed to analyse possible influence of sea lice Lepeophtheirus salmonis on marine Atlantic salmon Salmo salar survival. During the years 1992–2004, trawling for wild migrating post-smolts was performed in Trondheimsfjord, a fjord in which no Atlantic salmon aquaculture activity is permitted. Prevalence and intensity of sea lice infections on migrating wild post-smolts differed between years. A correlation analysis between 1 sea-winter (SW) Atlantic salmon catch statistics from the River Orkla (a Trondheimsfjord river) and sea lice infections on the migrating smolts in the Trondheimsfjord was not significant. Up to 2% reduction in adult returns due to sea-lice infection was expected. In addition, experimental releases from 1996 to 1998 with individually tagged groups of hatchery-reared Atlantic salmon smolts given protection against sea-lice infection was performed. Higher recaptures of adult Atlantic salmon from 1998 treated smolts compared to the control group may correspond to high abundance of sea lice found on the wild smolt, and may indicate influence on post-smolt mortality. These studies indicate that post-smolt mortality in Trondheimsfjord is marginally influenced by sea lice infection; however, the methods for assessing wild smolt mortality might be insufficient. Higher infections of sea lice farther out in the fjord may indicate more loss in Atlantic salmon returns in some years.     

Sea louse infection of juvenile sockeye salmon in relation to marine salmon farms on Canada's west coast

Background

Pathogens are growing threats to wildlife. The rapid growth of marine salmon farms over the past two decades has increased host abundance for pathogenic sea lice in coastal waters, and wild juvenile salmon swimming past farms are frequently infected with lice. Here we report the first investigation of the potential role of salmon farms in transmitting sea lice to juvenile sockeye salmon (Oncorhynchus nerka).

Methodology/Principal Findings

We used genetic analyses to determine the origin of sockeye from Canada''s two most important salmon rivers, the Fraser and Skeena; Fraser sockeye migrate through a region with salmon farms, and Skeena sockeye do not. We compared lice levels between Fraser and Skeena juvenile sockeye, and within the salmon farm region we compared lice levels on wild fish either before or after migration past farms. We matched the latter data on wild juveniles with sea lice data concurrently gathered on farms. Fraser River sockeye migrating through a region with salmon farms hosted an order of magnitude more sea lice than Skeena River populations, where there are no farms. Lice abundances on juvenile sockeye in the salmon farm region were substantially higher downstream of farms than upstream of farms for the two common species of lice: Caligus clemensi and Lepeophtheirus salmonis, and changes in their proportions between two years matched changes on the fish farms. Mixed-effects models show that position relative to salmon farms best explained C. clemensi abundance on sockeye, while migration year combined with position relative to salmon farms and temperature was one of two top models to explain L. salmonis abundance.

Conclusions/Significance

This is the first study to demonstrate a potential role of salmon farms in sea lice transmission to juvenile sockeye salmon during their critical early marine migration. Moreover, it demonstrates a major migration corridor past farms for sockeye that originated in the Fraser River, a complex of populations that are the subject of conservation concern.     

Sex‐selective Predation by Threespine Sticklebacks on Sea Lice: A Novel Cleaning Behaviour

Cleaning interactions have been described in a wide range of fish species and other taxa. We observed a novel cleaning behaviour during a study of the transmission dynamics of sea lice (Lepeophtheirus salmonis) between juvenile pink salmon (Oncorhynchus gorbuscha) and threespine sticklebacks (Gasterosteus aculeatus) in the Broughton Archipelago, British Columbia, Canada. Experiments showed that sticklebacks significantly reduced the number of sea lice on individual juvenile salmon. Adult female lice were preferentially consumed by sticklebacks, and gravid female lice also experienced egg string cropping. Overall, 76% of gravid female lice experienced either consumption, egg string cropping, or both by sticklebacks. This preference by sticklebacks for female parasites may stem from female lice being larger than males and the added nutritional value of egg strings on gravid females. Cleaning by sticklebacks can potentially have an impact on sea louse populations on wild juvenile salmon.     

Spatial and temporal variations in sea lice (Copepoda: Caligidae) infestations of three salmonid species farmed in net pens in southern Chile

Sea lice infestations have become a major health problem for farmed salmonids throughout the world including Chile. In southern Chile, 6 geographical areas, divided into 22 geographical zones with a total of 127 salmon farming centers and 1519 sea pens, were regularly sampled from December 1999 to April 2002. A linear mixed-effects model (LME) approach was used to describe the infestations of adult forms of sea lice on 3 salmonid species farmed in southern Chile. The variables fish species, water temperature, water salinity, fish weight, juvenile parasite count, pen shape, treatment status in previous month and the interaction of previous and current month treatments were found to be statistically significant fixed effects for the population sampled. The most susceptible species to sea lice infestation was rainbow trout Oncorhynchus mykiss, while the least susceptible species was coho salmon O. kisutch. Fishes in pens treated in the previous month with avermectins were associated with the smallest sea lice count compared to fishes in pens not treated or treated with other products. The variability in sea lice infestations in areas and zones within areas was not statistically significant when controlling for the previously mentioned fixed variables. The variability between centers, the within-pen variability, and the interaction between within-pen effect and the date of measurement were statistically significant and not explained by the fixed effects. Potential sources for this variability are discussed. We conclude that the epidemiology of sea lice infestations in farmed salmonids in southern Chile is complex and in need of further study.     

Temporal and spatial patterns of sea lice levels on sea trout in western Scotland in relation to fish farm production cycles

The relationship between aquaculture and infestations of sea lice on wild sea trout (Salmo trutta) populations is controversial. Although some authors have concluded that there is a link between aquaculture and lice burdens on wild fish, others have questioned this interpretation. Lice levels have been shown to be generally higher on Atlantic salmon farms during the second years of two-year production cycles. Here we investigate whether this pattern relates to lice burdens on wild fish across broad temporal and spatial axes. Within Loch Shieldaig across five successive farm cycles from 2000 to 2009, the percentage of sea trout with lice, and those above a critical level, were significantly higher in the second year of a two-year production cycle. These patterns were mirrored in 2002–2003 across the Scottish west coast. The results suggest a link between Atlantic salmon farms and sea lice burdens on sea trout in the west of Scotland.     

The potential of carbaryl as a treatment for sea lice infestations of farmed Atlantic salmon, Salmo salar L.

The treatment of sea lice, Lepeophtheirus salmonis infestations on farmed Atlantic salmon, Salmo salar requires regular bathing of the fish in diachlorvos (DDVP), 2,2-dichlorovinyl dimethyl phosphate, with subsequent release to the marine environment of spent pesticide. The toxicity of a possible alternative compound, carbaryl, l-naphthyl N-methylcarbamate) to kill sea lice and Atlantic salmon was examined. Preliminary studies with carbaryl indicate the potential advantage of a reduced treatment dose was outweighed by the greater persistence of both the parent compound and of its toxic degradation product. Hence this carbamate insecticide is unlikely to be considered by regulatory authorities as an alternative treatment for sea lice infestations.     

Transmission dynamics of parasitic sea lice from farm to wild salmon

Marine salmon farming has been correlated with parasitic sea lice infestations and concurrent declines of wild salmonids. Here, we report a quantitative analysis of how a single salmon farm altered the natural transmission dynamics of sea lice to juvenile Pacific salmon. We studied infections of sea lice (Lepeophtheirus salmonis and Caligus clemensi) on juvenile pink salmon (Oncorhynchus gorbuscha) and chum salmon (Oncorhynchus keta) as they passed an isolated salmon farm during their seaward migration down two long and narrow corridors. Our calculations suggest the infection pressure imposed by the farm was four orders of magnitude greater than ambient levels, resulting in a maximum infection pressure near the farm that was 73 times greater than ambient levels and exceeded ambient levels for 30 km along the two wild salmon migration corridors. The farm-produced cohort of lice parasitizing the wild juvenile hosts reached reproductive maturity and produced a second generation of lice that re-infected the juvenile salmon. This raises the infection pressure from the farm by an additional order of magnitude, with a composite infection pressure that exceeds ambient levels for 75 km of the two migration routes. Amplified sea lice infestations due to salmon farms are a potential limiting factor to wild salmonid conservation.     

Temporal stability of sea louse Lepeophtheirus salmonis Krøyer populations on Atlantic salmon Salmo salar L. of wild, farm and hybrid parentage

Atlantic salmon salmo salar smolts of wild, hybrid and farmed parentage were individually tagged then reared in a sea cage for 8 months. The fish were sampled three times during this period. On all occasions, farmed Atlantic salmon displayed the highest abundance and density of sea lice Lepeophtheirus salmonis , whilst no significant differences were observed between hybrid and wild Atlantic salmon. Percentage variation between the lowest and highest infected groups was as high as 175 and 144% for L. salmonis abundance and density respectively (sample 2). The temporal stability of interindividual sea lice infection levels was investigated pair‐wise between samples using correlation (sample 1 v . 2, 1 v . 3 and 2 v . 3). When calculated using sea louse abundance, correlations ranged from r ² = 0·11, P  < 0·01 to r ² = 0·39, P  < 0·001, but, when the effects of fish size were controlled for by converting abundance to density, all correlations were <  r ² = 0·1. Therefore, these data indicate that a fish's relative infection level in one sample was a weak predictor of its relative infection level in another sample. This suggests that identification of individual Atlantic salmon that display reduced susceptibility to sea lice, may be problematic.     

Tidal and diel timing of river entry by adult Atlantic salmon returning to the Aberdeenshire Dee, Scotland

The timing of salmon migration from the estuary of the Aberdeenshire Dee into the river in relation to tidal phase and time of day was studied by combined acoustic and radio-tracking of individual fish and by analysing records of untagged fish from a resistivity fish counter 0.8 km upstream from the tidal limit. Up-estuary movements that led to river entry were predominantly nocturnal and tended to occur during the ebb tide. Penetration into the non-tidal reaches of the river also tended to occur at night, but the timing of salmon movements was no longer significantly associated with tidal phase. The tracking data suggested that the reduction in the strength of the association between salmon migration and tidal phase resulted from variability in rates of progress from the estuary to the river. This variability may have been random, or related to changes in migratory behaviour during entry to fresh water. A simulation of upstream progress by groups of salmon illustrated the decay of the relationship between salmon movements and tidal phase and the advance of the average tidal phase of observed salmon movements with increasing distance of the observation point from where movements were initiated. The magnitude of these effects depended on the average rate of upstream progress and variability in the rate of progress, being greatest when upstream progress was slow and variable. These results highlight a limitation of point observations of migration with regard to identifying environmental stimuli for migration and quantifying their effects.     

Changes in Atlantic salmon (Salmo salar) epidermal mucus protein composition profiles following infection with sea lice (Lepeophtheirus salmonis)

The mucus protein profile of Atlantic salmon (Salmo salar) and changes due to infection with sea lice (Lepeophtheirus salmonis) were examined. Two-dimensional gel electrophoresis was performed on salmon skin mucus and comparisons between control and infected fish mucus were made. LC MS/MS identified intracellular proteins, calmodulin, actin, and hemopexin and plasma proteins, such as apolipoproteins, lectin, plasminogen and transferrin. Plasma proteins in the mucus may result from either direct expression by epidermal cells, leakage of plasma or via a secondary circulation system. Therefore, RT-PCR was used to measure mRNA of transferrin and lectin in Atlantic salmon skin. Transferrin expression was observed suggesting direct expression by the epidermis. Lectin expression was not detected suggesting another mechanism of entry into mucus, either leakage from plasma or secondary circulation. The lack of observable albumin on 2D gels, suggests that mucus lectin may arise from the secondary circulation route. Interestingly, β-actin was a significant component of Atlantic salmon mucus. Cleaved actin and transferrin fragments were observed and positively correlated with sea lice infection suggestive of proteolytic activity. Increased levels of cleaved transferrin during sea lice infection may activate the nitrous oxide response of salmon macrophages, as part of the fish's immune response to sea lice infection.     

Lessons from sea louse and salmon epidemiology

Effective disease management can benefit from mathematical models that identify drivers of epidemiological change and guide decision-making. This is well illustrated in the host–parasite system of sea lice and salmon, which has been modelled extensively due to the economic costs associated with sea louse infections on salmon farms and the conservation concerns associated with sea louse infections on wild salmon. Consequently, a rich modelling literature devoted to sea louse and salmon epidemiology has been developed. We provide a synthesis of the mathematical and statistical models that have been used to study the epidemiology of sea lice and salmon. These studies span both conceptual and tactical models to quantify the effects of infections on host populations and communities, describe and predict patterns of transmission and dispersal, and guide evidence-based management of wild and farmed salmon. As aquaculture production continues to increase, advances made in modelling sea louse and salmon epidemiology should inform the sustainable management of marine resources.     

Enzymes released from Lepeophtheirus salmonis in response to mucus from different salmonids

Adult and mobile preadult sea lice Lepophtheirus salmonis were incubated with mucus samples from rainbow trout (Oncorhynchus mykiss), coho salmon (O. kisutch), Atlantic salmon (Salmo salar), and winter flounder (Pseudopleuronectes americanus) to determine the response of L. salmonis to fish skin mucus as assessed by the release of proteases and alkaline phosphatase. There was variation in the release of respective enzymes by sea lice in response to different fish. As well, sealice collected from British Columbia responded differently than New Brunswick sea lice to coho salmon mucus. Fish mucus and seawater samples were also analyzed using protease gel zymography to observe changes in the presence of low molecular weight (LMW) proteases after L. salmonis incubation. Significantly higher proportions of sea lice secreted multiple bands of L. salmonis-derived LMW proteases after incubation with rainbow trout or Atlantic salmon mucus in comparison with seawater, coho salmon, or winter flounder mucus. Susceptibility to L. salmonis infections may be related to the stimulation of LMW proteases from L. salmonis by fish mucus. The resistance of coho salmon to L. salmonis infection may be due to agents in their mucus that block the secretion of these LMW proteases or factors may exist in the mucus of susceptible species that stimulate their release.     

Notes on the behaviour of lumpfish in sea pens with and without Atlantic salmon present

The behaviour of lumpfish, Cyclopterus lumpus L., in sea pens, with and without Atlantic salmon, Salmo salar L., present, was assessed by underwater camera technology. Behaviour was classified by recording the principal activity of individual fish for 30-s intervals. The majority of daylight time was spent actively foraging for food. Antagonistic behaviour between Atlantic salmon and lumpfish was not observed during the whole experimental period and no mortality was seen in either species. Cleaning behaviour, but at low frequency, was observed as lumpfish cleaned sea lice off Atlantic salmon. Significantly lower sea lice infection levels were seen on Atlantic salmon when reared together with lumpfish compared to the control group without lumpfish. Feeding behaviour can be classified as strongly opportunistic.     

Sea lice escape predation on their host

Parasites seldom have predators but often fall victim to those of their hosts. How parasites respond to host predation can have important consequences for both hosts and parasites, though empirical investigations are rare. The exposure of wild juvenile salmon to sea lice (Lepeophtheirus salmonis) from salmon farms allowed us to study a novel ecological interaction: the response of sea lice to predation on their juvenile pink and chum salmon hosts by two salmonid predators-coho smolts and cut-throat trout. In approximately 70% of trials in which a predator consumed a parasitized prey, lice escaped predation by swimming or moving directly onto the predator. This trophic transmission is strongly male biased, probably because behaviour and morphology constrain female movement and transmission. These findings highlight the potential for sea lice to be transmitted up marine food webs in areas of intensive salmon aquaculture, with implications for louse population dynamics and predatory salmonid health.     

Genetic differentiation of populations of the copepod sea louse Lepeophtheirus salmonis (Krøyer) ectoparasitic on wild and farmed salmonids around the coasts of Scotland: Evidence from RAPD markers

Sea trout are the sea-going migratory form of the freshwater brown trout (Salmo trutta L.) and since 1989 there have been marked declines in their stocks on the west coasts both of Scotland and Ireland. Various factors have been attributed as possible causal agents in these stock declines, including fresh water acidification, overfishing, climatic fluctuations, habitat degradation and sea lice parasitic burdens. The putative impact of infestations of sea trout by the ectoparasitic copepod sea louse, Lepeophtheirus salmonis (Krøyer), has featured prominently in the controversy, especially with regard to the role of inshore commercial salmon farms as a possible source of infestation of wild salmonids by sea lice. This study focused on the population genetics of L. salmonis around the coasts of Scotland: We sampled fish from wild and cultured stocks and included salmon (Salmo salar L.), rainbow trout (Oncorhynchus mykiss Walbaum) and sea trout as host species. Analyses of allozyme variation of sea lice were confined to data for two polymorphic loci (Fum, Got-2) and conformed to our initial expectation — that the inclusion of a planktonic larval phase in the life cycle of the copepod, in addition to the high mobility of the host fish, would enhance gene flow and preclude genetic differentiation of L. salmonis populations as a result of random drift alone. DNA polymorphism was quantified by means of PCR and RAPD analysis. Six primers were screened for 16 samples (from wild and farmed salmon, wild sea trout and farmed rainbow trout) — including the east, north and west coasts of Scotland — and the data analyzed by AMOVA (Analysis of Molecular Variance). In contrast to the allozyme results, the RAPD analysis showed striking patterns of genetic differentiation around the coasts of Scotland. The overall pattern was one of genetic homogeneity of L. salmonis populations sampled from wild salmon and sea trout. All of the L. salmonis samples taken from farmed salmon and rainbow trout did, however, show highly significant levels of genetic differentiation, both between wild and farmed salmonids and among the various farms themselves. Evidence of high levels of small-scale spatial or temporal heterogeneity of RAPD marker band frequencies was shown for the one farm from which repeat samples (July and November, 1995) were analysed. Samples of sea lice taken from west coast wild sea trout subjected to RAPD analysis also revealed the occurrence of putative “farm markers” in some individual parasites, indicating that they had possibly originated from salmon farms.     

Seasonal occurrence of sea lice Lepeophtheirus salmonis on sea trout in two north Norwegian fjords

Seasonal occurrence of the parasitic copepod Lepeophtheirus salmonis (sea lice) was studied from March to December 2001 in two large north Norwegian sill fjords without fish farming activity, the Ranafjord and the Balsfjord. Anadromous brown trout Salmo trutta (sea trout) in both fjords had a low infestation rate during all sampling periods, but followed a seasonal pattern. During early and late winter (November to December and March to April) and spring (May to June), the prevalence varied from 0 to 25% and the abundance was <0·5 sea lice. Adults dominated (92%) during this period, particularly gravid females. In both fjords, the highest prevalence was during September (80–81%, all stages represented). In Ranafjord, the abundance and mean intensity during this month was 6·8 and 8·6 sea lice, respectively, while in Balsfjord it was 3·6 and 4·5 sea lice, respectively. Fish were captured at temperatures down to 1° C and at full strength sea water which is supposed to cause osmoregulatory problems for the fish. This observation has implications for the understanding of high‐latitude sea trout behaviour and can also make the fish more vulnerable to heavy sea lice infestation during this period. It is suggested that winter running sea trout help to maintain a self replicating local population of sea lice within such fjord systems where other possible hosts ( e.g . farmed Atlantic salmon Salmo salar ) are not present during a whole year cycle.