A comparison of epidemiological patterns of salmon lice, Lepeophtheirus salmonis, infections on farmed Atlantic salmon, Salmo salar L., in Norway and Scotland

This paper examines two large national data sets collected over several years and contrasts the patterns of sea lice, Lepeophtheirus salmonis (Krøyer, 1837), infections, the use of treatments and the occurrence of chalimus peaks between Norwegian and Scottish farms. Infection levels in Scotland were significantly higher in general over the period under study. For the chalimus stage group in the first quarter of the year, Norwegian mean abundance stayed below 10 lice per m² while Scottish means reached 45 lice per m² of fish skin per m³ of water. Both countries had more chalimus in summer than at other times of year, but in the last 3 months of the year Scottish fish had, on average, two to four times as many chalimus as Norwegian fish. Peaks of chalimus abundance were more frequent in Scotland, particularly in winter, but the most prominent peaks occurred in summer in both countries. In Scotland a marked mid-year build-up of mobile pre-adult and adult stages was seen, and both countries showed a tendency for mobile counts on the second year fish to increase towards the end of the year. Scottish fish carried, on average, three times as many mobile lice per m² of skin as Norwegian fish in the last 3 months of the year. The difference in lice loads was reflected in the greater use of veterinary medicines on Scottish farms. The higher infection levels in Scotland may be due to shallower and more enclosed water bodies used for farming, smaller and shallower pens, differences in sea water temperatures or in access to appropriate medication. The results highlight the importance of ensuring that effective veterinary medicines are available in the UK for the control of infection.     

Differences in Lepeophtheirus salmonis abundance levels on Atlantic salmon farms in the Broughton Archipelago, British Columbia, Canada

Sea lice data collected from Atlantic salmon farms in the Broughton Archipelago between 2003 and 2005 were examined for inter-regional differences in mobile Lepeophtheirus salmonis (Krøyer) abundance using the generalized linear model procedure. Factors such as age of the salmon populations, location of farms and time of year had a significant effect on the abundance of the mobile stages of L. salmonis whereas water temperature and salinity did not. Separate evaluation of SLICE® treatment data found no significant difference in treatment frequency among the areas but did show that there were significantly lower numbers of farm treatments during the summer months when compared with other seasons. The role of environment and wild fish in influencing sea lice abundance on the farmed salmon is discussed. The findings suggest that effective management programmes for sea lice should not only be based on geographical location but should take into account other factors which could influence lice abundance levels.     

Variation in sensitivity of sea lice [Lepeophtheirus salmonis (Krøyer)] to dichlorvos on Irish salmon farms in 1991–92

Variation in sensitivity of Lepeophtheirus salmonis (Krøyer) to dichlorvos in static toxicity tests was recorded on salmon farms in Ireland. Five hour LC50s ranged from 60 to 202 μg l ^?1 at 15 °C for five farms in Autumn 1991 and 76–151 μg l^?1 at 10 °C for four farms in Spring of 1992. Resistance ratios, calculated using the most sensitive sites as baseline, were low and ranged from 1.32 to 3.27 in 1991 and 1.28–1.98 in 1992. Repeat sampling on three farms in 1991 and 1992 also indicated short‐term changes in sensitivity to dichlorvos within farms.     

Changes in epidemiological patterns of sea lice infestation on farmed Atlantic salmon, Salmo salar L., in Scotland between 1996 and 2006

Analyses of a unique database containing sea lice records over an 11 year period provide evidence of changing infestation patterns in Scotland. The data, collected from more than 50 commercial Atlantic salmon farms, indicate that both species of sea lice commonly found in Scotland, Lepeophtheirus salmonis and Caligus elongatus, have declined on farms over the past decade. Reductions for both species have been particularly marked since 2001 when more effective veterinary medicines became available. Treatment data were also available in the database and these show a growing trend towards the use of the in-feed medication emamectin benzoate (Slice), particularly in the first year of the salmon production cycle. However, this trend towards single product use has not been sustained in 2006, the latest year for which data are available. There is some evidence of region to region variation within Scotland with the Western Isles experiencing higher levels of infestation. However, compared to the levels observed between 1996 and 2000, all regions have benefited from reduced lice infestation, with the overall pattern showing a particular reduction in the second and third quarters of the second year of production.     

Sea lice treatments, management practices and sea lice sampling methods on Atlantic salmon farms in the Bay of Fundy, New Brunswick, Canada

Proper monitoring of all stages of sea lice is imperative to ensure the strategic timing of treatments. Although sea lice receive close attention and management for production purposes, there are no regulations for the reporting of lice burdens on salmon farms in Atlantic Canada, nor are there officially standardized protocols for conducting sea lice counts in the field. The purpose of this study was to complete a survey of Atlantic salmon farms in the Bay of Fundy, NB, Canada. The survey addressed many aspects of sea lice monitoring and control, including methods for sea lice sampling at the site, types of treatments administered for control of sea lice and management practices and decisions regarding the control of sea lice. A total of 83 surveys were completed from July to December 2002. Twelve of the 60 respondents owned and/or operated more than one site and completed a survey for each of their sites at the time of the interview. The position of individuals surveyed included 48 site managers (representing 63 sites), three site owners (representing 11 sites), eight individuals who were both owner and manager (representing nine sites) and one site veterinarian (representing one site). Surveys were completed on 30 sites from the Limekiln Bay and surrounding areas, 23 sites from Grand Manan Island, 18 sites from Deer Island, six sites from Campobello Island and six sites from Passamaquoddy Bay. Fish farmers, in concert with their veterinarians, generally perform intensive monitoring and control practices. However, there is a continued reliance on emamectin benzoate (SLICE^®) for sea lice control on Atlantic salmon farms in the Bay of Fundy, which raises concerns regarding the potential for sea lice to develop resistance to the drug.     

An assessment of salmon farms and wild salmonids as sources of Lepeophtheirus salmonis (Krøyer) copepodids in the water column in Loch Torridon, Scotland

Wild salmonids and farmed salmon can both be sources of Lepeophtheirus salmonis (Krøyer, 1838) larvae . Farmed salmon smolts free of L. salmonis infections are stocked in sea cages and may subsequently contract L. salmonis infections, probably from wild fish. The contribution of gravid L. salmonis at Atlantic salmon, Salmo salar L., farms to populations of L. salmonis larvae in the water column has in the past been based on estimated parameters, such as louse fecundity. This present study augments these calculations by combining empirical data on densities of infective L. salmonis copepodids in the field with estimates of the number of gravid L. salmonis on farmed and wild salmonids in Loch Torridon. Data collected between 2002 and 2007 show a significant correlation between mean densities of L. salmonis copepodids recovered in the water column and the numbers of gravid L. salmonis at the local salmon farms. Generally, the farms with greatest numbers of salmon were observed to have stronger correlations with densities of copepodids in the water than the farms with fewer fish. The study suggests that louse management approaches, e.g. treatment trigger levels, need to take account of individual farm biomass, or numbers of fish. This study highlights the importance of control of L. salmonis on salmon farms for the co-existence of both wild salmonid populations and the aquaculture industry.     

Sea lice, Lepeophtheirus salmonis, transfer between wild sympatric adult and juvenile salmon on the north coast of British Columbia, Canada

We examine sea lice, Lepeophtheirus salmonis , on juvenile and adult salmon from the north coast of British Columbia between 2004 and 2006 in an area that does not at present contain salmon farms. There is a pronounced zonation in the abundance of L. salmonis on juvenile pink salmon, Oncorhynchus gorbuscha , in the Skeena and Nass estuaries. Abundances in the proximal and distal zones of these estuaries are 0.01 and 0.05 respectively. The outer zones serve as feeding and staging areas for the pink salmon smolts. Returning Chinook, Oncorhynchus tshawytscha , and coho salmon, Oncorhynchus kisutch , concentrate in these areas. We collected data in 2006 to examine whether L. salmonis on returning adult salmon are an important source of the sea lice that appear on juvenile pink salmon. Nearly all (99%) of the sea lice on returning Chinook and over 80% on coho salmon were L. salmonis. Most of the L. salmonis were motile stages including many ovigerous females. There was a sharp increase in the abundance of sea lice on juvenile pink salmon smolts between May and July 2006 near the sites of adult captures. As there are no salmon farms on the north coast, few sticklebacks, Gasterosteus aculeatus , and very few resident salmonids until later in the summer, it seems that the most important reservoir of L. salmonis under natural conditions is returning adult salmon. This natural source of sea lice results in levels of abundance that are one or two orders of magnitude lower than those observed on juvenile pink salmon in areas with salmon farms such as the Broughton Archipelago.     

Distributions of planktonic sea lice larvae Lepeophtheirus salmonis in the inter-tidal zone in Loch Torridon, Western Scotland in relation to salmon farm production cycles

This study examines the relative density of sea lice, Lepeophtheirus salmonis (Krøyer), larvae in the inter‐tidal areas of Loch Torridon, a fjordic sea loch in Western Scotland and the site of several commercial salmon farms. Samples of planktonic sea lice were collected in each spring from 1999 to 2003 near the mouth of the River Shieldaig, where sea trout have been shown to congregate in the first few days after going to sea, and from 2001 to 2003 near the mouth of the River Balgy. No larval sea lice were found near the mouth of the River Shieldaig in spring 2000 or 2002, when gravid female sea lice were absent on local fish farms. The same finding was observed in 2002 at the mouth of the River Balgy. In springs of 1999, 2001 and 2003, peak densities of larval sea lice of 33, 94 and 143 m^?3, respectively, were observed at the mouth of the river Shieldaig. At the mouth of the River Balgy, peak densities of 74 and 78 lice m^?3 were observed in 2001 and 2003 respectively. At the time these samples were collected, the two local salmon farms were in the second year of their production cycle and gravid female sea lice were present. Samples of larval sea lice were also obtained year round, at approximately weekly intervals, from a 50‐m transect at the mouth of the River Shieldaig, from March 2001 to June 2003, and compared with frequencies of gravid female sea lice on the two local salmon farms. High levels were found in the winter. No planktonic sea lice were found in this transect when gravid females were not present on the local fish farms.     

A mathematical model of the growth of sea lice, Lepeophtheirus salmonis, populations on farmed Atlantic salmon, Salmo salar L., in Scotland and its use in the assessment of treatment strategies

Sea lice are a persistent problem for farmed and wild salmonid populations. Control can be achieved through the use of veterinary medicines. A model was developed to describe the patterns of sea lice infection on salmon farms in Scotland and to predict the likely effect of various treatment strategies. This model takes into account development rates and mortality using compartments representing life history stages and external infection pressure. The national sea lice infection pattern was described using parameters representing stage survival, background infection levels and egg viability rates. The patterns observed across farms varied greatly and the model gave broad agreement to observed trends with different parameters being required in the model for sites using hydrogen peroxide and cypermethrin treatments. The parameter estimates suggest that the background infection pressure on sites where cypermethrin was administered was higher than for those using hydrogen peroxide. Both models had comparable magnitudes of sensitivity with survival from one stage to another being the most sensitive parameter, followed by feedback rates at which gravid females produce eggs, with background infection levels the least sensitive. The effect of different cypermethrin treatment strategies was assessed using the model. Increasing treatments in a production cycle gave more effective control. However, the model showed that timing of treatments is most important if sea lice are to be effectively controlled.     

Resistance of sea lice, Lepeophtheirus salmonis (Krøyer), to hydrogen peroxide on farmed Atlantic salmon, Salmo salar L.

Hydrogen peroxide has been the only medicine used to treat salmon infected with sea lice [Lepeophtheirus salmonis (Krøyer)] on many farms in Scotland since 1992 and reports have suggested reduced treatment efficacy. The present study tests the sensitivity of sea lice to exposure to hydrogen peroxide under farm conditions and also in vivo by comparing lice from a farm with suspected resistance and lice from a farm that had not been pre‐exposed to hydrogen peroxide. In bin treatments on fish from a farm treated with hydrogen peroxide on 41 occasions, numbers of ovigerous lice declined by only 15% and 16% in two replicates and other mobile stages by 25%. Where hydrogen peroxide had not been used, ovigerous females declined by 90% and 87% and other mobile lice stages by 97% and 99%. These trials and observed poor efficacy of cage treatments, for example only 7.5% reduction in lice numbers when fish were treated with 2000 p.p.m. for 20 mins, indicated resistance of lice to hydrogen peroxide. The mechanisms involved in the development of resistance, possible genetic selection for lice with reduced carapace permeability or detoxifying enzymes such as catalase, or tolerance through induction by subtherapeutic exposure are reviewed. Implications for lice control strategies relying on hydrogen peroxide are discussed.     

Impact of co-infection with Lepeophtheirus salmonis and Moritella viscosa on inflammatory and immune responses of Atlantic salmon (Salmo salar)

This study was conducted to determine the effects of a co-infection with Moritella viscosa at different exposure levels of sea lice Lepeophtheirus salmonis in Atlantic salmon (Salmo salar). M. viscosa (1.14 × 10⁶ cfu/ml) was introduced to all experimental tanks at 10 days post-lice infection (dpLs). Mean lice counts decreased over time in both the medium lice co-infection (31.5 ± 19.0 at 7 dpLs; 16.9 ± 9.3 at 46 dpLs) and high lice co-infection (62.0 ± 10.8 at 7 dpLs; 37.6 ± 11.3 at 46 dpLs). There were significantly higher mortalities and more severe skin lesions in the high lice co-infected group compared to medium lice co-infected group or M. viscosa-only infection. Quantitative gene expression analysis detected a significant upregulation of genes in skin from the high lice co-infection group consistent with severe inflammation (il-8, mmp-9, hep, saa). Skin lesions retrieved throughout the study were positive for M. viscosa growth, but these were rarely located in regions associated with lice. These results suggest that while M. viscosa infection itself may induce skin lesion development in salmon, co-infection with high numbers of lice can enhance this impact and significantly reduce the ability of these lesions to resolve, resulting in increased mortality.     

Vertical movements of Atlantic salmon post-smolts relative to measures of salinity and water temperature during the first phase of the marine migration

Abstract  The migratory behaviour of hatchery-reared Atlantic salmon, Salmo salar L., post-smolts during the first phase of the marine migration was examined to assess their susceptibility to salmon lice, Lepeophtheirus salmonis (Krøyer), infestations. Swimming depths of eight post-smolts relative to the measured salinity and temperature were monitored for an average of 11.4 h following release outside the mouth of the River Eio using depth sensitive acoustic transmitters. Vertical salinity and temperature distributions were simultaneously recorded along the migratory route. Mean swimming depth was 1.7 m (individual mean 0.5–2.1 m). There was no overall preference among all the post-smolts for specific salinity concentrations. Typically post-smolts migrated the majority (68%) of their time at salinities <20 psu (brackish water), and as a result outside the reported salinity tolerances of sea lice. Furthermore, post-smolts chose the warmest water layer during their coastal migration.     

Efficacy of emamectin benzoate against sea lice infestations of Atlantic salmon, Salmo salar L.: evaluation in the absence of an untreated contemporary control

The efficacy of emamectin benzoate (SLICE) against sea lice infestations of Atlantic salmon, Salmo salar L., is typically assessed using untreated fish, or fish treated with alternative therapeutants, as controls. The State of Maine, USA, is currently under active management for the OIE-notifiable pathogen, infectious salmon anaemia virus (ISAV); consequently, neither control group is feasible in this region. Untreated salmon risk extensive damage from the ectoparasites, and threaten to increase vector-borne exposure or susceptibility of farms to ISAV; and the only treatment presently available in Maine is SLICE. However, because sea lice infestations are unlikely to resolve spontaneously, and response to treatment occurs within weeks, use of a pretreatment baseline is a reasonable alternative for confirmatory studies. We evaluated SLICE efficacy on Atlantic salmon farms in Cobscook Bay 2002-2005, in the absence of untreated controls, using pretreatment lice loads as a reference for calculation. Maximum efficacy ranged from 68% to 100% reduction from initial levels. Time-to-maximum efficacy ranged from 1 to 8 weeks after treatment initiation. Efficacy duration, measured between first reduction and first progressive rise in counts, ranged from 4 to 16 weeks.     

Safety and efficacy of emamectin benzoate administered in-feed to Atlantic salmon, Salmo salar L., smolts in freshwater, as a preventative treatment against infestations of sea lice, Lepeophtheirus salmonis (Krøyer)

The safety and efficacy of emamectin benzoate, administered in-feed to Atlantic salmon smolts, Salmo salar L., held in freshwater, was evaluated as a preventative treatment against sea lice, Lepeophtheirus salmonis, following transfer of fish to seawater.

In the safety study, salmon smolts held in freshwater were fed with diets containing emamectin benzoate at nominal doses of 0 (control), 50 (recommended dose) and 250 (5× recommended dose) μg kg⁻¹ fish day⁻¹ for 7 days (days 0–6). Actual dose rates, based on measured concentrations of emamectin benzoate in feed, differences in fish weight, and feed consumed, were 0, 54, and 272 μg kg⁻¹ day⁻¹, respectively. On day 9, fish were transferred to seawater and observed for 14 days. No differences in feeding response, coordination, behaviour, gross and histological appearance were observed between control fish and those that received 54 μg kg⁻¹ day⁻¹. Among smolts that received 272 μg kg⁻¹ day⁻¹, approximately 50% exhibited darker coloration, and one fish (1%) exhibited uncoordinated swimming behaviour. No pathognomonic signs of emamectin benzoate toxicity were identified.

In the efficacy study, smolts held in freshwater were fed an unmedicated ration (control group) or emamectin benzoate at 50 μg kg⁻¹ day⁻¹ (treated group) for 7 days (days 0–6). On day 9, fish were re-distributed to eight seawater tanks, each holding 30 control and 30 treated fish. On days 28, 56, 77 and 109, respectively, control and treated fish in two tanks were challenged with L. salmonis copepodites. When lice in each group reached chalimus stage IV, fish were sampled and the numbers of lice were recorded. Fish challenged at day 109 were sampled for the second time when lice were at the adult stage. Efficacy was calculated as the reduction in the mean number of lice on treated fish relative to the mean on control fish. Treatment with emamectin benzoate resulted in an efficacy of 85.0–99.8% in fish challenged at days 28–77, from the start of treatment, and lice counts were significantly lower (P<0.001) on treated fish than on controls. When fish challenged at day 109 were sampled at day 128, efficacy was 44.3%, but survival of chalimus to adult lice on treated fish was lower, and at day 159, efficacy had increased to 73%. These results demonstrate that treatment of salmon smolts with emamectin benzoate in freshwater was well tolerated and highly effective in preventing sea lice infestation following transfer of fish to seawater.     

The effect of light on the settlement of the salmon louse, Lepeophtheirus salmonis, on Atlantic salmon, Salmo salar L

The salmon louse, Lepeophtheirus salmonis, is an ectoparasitic copepod that infests both wild and farmed salmonid fish. Salmon lice are a major disease problem in the farming of Atlantic salmon, Salmo salar L., and the possibility of salmon lice playing a role in the decline of wild anadromous stocks has also been raised. Lepeophtheirus salmonis can detect a range of stimuli (pressure/moving water, chemicals and light) in the external environment. However, the response thresholds to various stimuli, and the spatial and temporal scales over which they operate in the context of host location, are largely unknown. In this context, we attempted to determine whether salmon lice copepodids settle onto hosts more effectively, or at different locations on the fish's body, under different qualities of light. Lice settlement trials were conducted under three lighting conditions; L1: unpolarized under ultraviolet A (UVA)-through visible; L2: unpolarized without UVA (control); L3: 100% linearly polarized without UVA. A dark control was also conducted. No statistically significant difference in lice settlement was found. While changes in light intensity are involved in host detection at spatial scales on the order of metres, the results presented here suggest that it is not the primary sensory modality underlying host location at smaller spatial scales (cm to mm).     

Early‐stage sea lice recruits on Atlantic salmon are freshwater sensitive

Sea lice are significant parasites of marine and brackish farmed fishes. Freshwater bathing is a potential control option against numerous sea lice species, although has been viewed as futile against those that are capable of tolerating freshwater for extended periods. By comparing freshwater survival times across host‐attached stages of Lepeophtheirus salmonis (Krøyer), a key parasite in Atlantic salmon farming, we show the first attached (copepodid) stage undergoes 96–100% mortality after 1 h in freshwater, whereas later attached stages can tolerate up to 8 days. Thus, regular freshwater bathing methods targeting the more susceptible attached copepodid stage may successfully treat against L. salmonis and potentially other sea lice on fish cultured in marine and brackish waters.     

The effect of artificial light treatment and depth on the infestation of the sea louse Lepeophtheirus salmonis on Atlantic salmon (Salmo salar L.) culture

Two field studies were carried out with farmed Atlantic salmon (Salmo salarL.) in sea cages to examine various effects of artificial light (AL) and the vertical distribution of salmon on lice infestation.

The use of AL light caused an overall increase in lice infestation in both experiments. The first study showed that salmon held at 0–4 m depth in cages developed higher infestation than salmon held at greater depths (4–8 and 8–12 m) under both natural light (NL) and AL. In the second study, salmon maintained in 14-m deep sea cages that were exposed to AL with different light intensities. The AL treatments resulted directly in different diel and seasonal patterns of vertical distribution of the salmon and also different temporal patterns in lice infestations. So indirectly the infestation pattern appeared to be correlated with median day-time swimming depth of the salmon.