Marine Aquaculture Site: Loch Alsh - Sepa
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Marine Aquaculture Site: Loch Alsh ANNEX 1 : SEA LICE MANAG EMENT AND EFFICACY STATEM ENT MARINE HARVEST ( SCOTLAND) LIMITED WRITTEN BY RUTH PATERSON JULY 2018; REVIEWED SEPTEMBER 2018 Registered in Scotland No. 138843 Stob Ban House, Glen Nevis Business Park, Registered Office, Fort William, PH33 6RX 1st Floor, Admiralty Park Ruth. Paterson@marineharvest. com Admiralty Road Rosyth Stob Ban House, Glen Nevis Business Park, FIFE Fort William, PH33 6RX KY11 2YW http:// marineharvest. com
2 | 16 Table of Contents Table of Contents ...........................................................................................................................................2 Table of Figures ..............................................................................................................................................2 1. Introduction ............................................................................................................................................ 3 1 Policy and Legislation Framework ........................................................................................................ 3 1.1 Code of Good Practice ................................................................................................................... 3 1.2 Aquaculture and Fisheries Scotland Act 2013 ............................................................................. 3 1.3 Current Compliance ...................................................................................................................... 4 2 Historic Sea Lice Challenges .................................................................................................................. 4 2.1 Marine Harvest Response to Rising Sea Lice Numbers ..............................................................5 3 Development of Lice Management Tools ..............................................................................................6 3.1 Cleaner Fish ................................................................................................................................... 6 3.2 Non-Medicinal Lice Treatments ................................................................................................... 7 3.2.1 Hydrolicer Units ....................................................................................................................... 7 3.2.2 Thermolicer Units ..................................................................................................................... 7 3.2.3 Freshwater Treatments ............................................................................................................ 9 3.2.4 Pen Modifications ................................................................................................................... 10 3.3 Medicinal Treatments ................................................................................................................. 10 3.4 Good Husbandry Practice ............................................................................................................11 4 Treatment Selection and Hierarchy .................................................................................................... 12 5 Conclusion ............................................................................................................................................12 Appendix 1 Example Treatment Plan: Loch Alsh ....................................................................................... 13 6 References ............................................................................................................................................. 16 Table of Figures Figure 1. Adult Female Lice Levels vs Marine Scotland Thresholds 2016-2018. ................... 4 Figure 2. Average monthly adult female levels 2014-2018. ................................................... 5 Figure 3. Marine Harvest cleaner fish deployment plan 2018-2022. ...................................... 6 Figure 4. Schematic of a thermolicer system used within Marine Harvest.............................. 8 Figure 5. Flusher results from 2016 to 2018 for adult female lice clearance levels. ............... 9
3 | 16 1. Introduction Effective sea lice management is a critical requirement to protect the health and welfare of farmed fish and to minimise the risk of potential impacts on wild salmonids. Marine Harvest Scotland) Ltd. has incrementally introduced a suite of internal policies, targets, and capital investment programmes1 to significantly improve sea lice management, in parallel with gradual reduction of medicinal treatments. The company is now equipped with an increased range and capacity of sea lice management tools, not previously available up until early 2016. This document summarises these changes, outlines the general efficacy of each treatment and illustrates how these can be applied in an integrated manner to effectively manage sea lice issues. In this document, Marine Harvest sets out its policies and aims on sea lice management and details the operational approach and resources to target sites to meet regulatory and industry guideline treatment thresholds for sea lice, but also importantly the stricter operational intervention thresholds the company apply and aspire to meet. This document outlines how the investment in sea lice management tools is being applied across Marine Harvest; the efficacy that is being achieved with these new tools; and the declining levels of parasitic loading that is now being seen as a result. 1 Policy and Legislation Framework 1.1 Code of Good Practice In 2006, the Code of Good Practice for Scottish Finfish Aquaculture (CoGP)2 was introduced covering the production of all types of finfish farmed in Scotland and the good practice stated within underpins all aspects of fish farming operations ( CoGP, 2003). The National Treatment Strategy for the Control of Sea Lice on Scottish Salmon Farms was subsequently revised and incorporated into the CoGP, taking new initiatives on maximising the efficacy of sea lice treatments and medicines, and minimising sea louse resistance. A primary objective of the revised strategy is the target of zero adult female lice on farmed fish in the spring period when wild salmonids are migrating, achieved by coordinated spring treatments, conducted by all farms within defined management areas that are in their second year of production, or if lice numbers are above the defined management threshold levels. These levels are as follows: 0.5 adult female lice per fish from February to June to minimise any potential for interaction with the wild smolt run; and One adult female per fish from July to January. 1.2 Aquaculture and Fisheries Scotland Act 2013 In 2007 the Scottish Parliament made provisions in relation to fish farms, specifically with regards to the control of sea lice (amended in 2013) which put into regulatory context minimum 1 In the last 2 years, Marine Harvest has invested over £ 16 million in the acquisition and development of the latest available, non-medicinal methods of sea lice control. These new non-medicinal treatments, only developed since late 2015 include Hydrolicer” systems, “ Thermolicer” systems, and new Freshwater treatment systems http://thecodeofgoodpra 2 http:// thecodeofgoodpractice. co.uk/ ctice.co.uk/
4 | 16 legal standards covering the health and welfare needs of farmed fish, including measures for the prevention, control and reduction of parasites on fish farms. In 2017 Marine Scotland significantly updated their guidance and monitoring to ensure that all sea lice control measures being implemented on Scottish fish farms can be considered as satisfactory. This new ( 2017) monitoring scheme includes a new mandatory reporting level where any farm reaching an average of 3.0 adult female lice per fish requires to be reported to the Fish Health Inspectorate ( FHI). If such a lice level is reached, increased monitoring by the FHI is implemented and continued until the adult female lice count is reduced to below the reporting level of 3.0. However, if a farm reaches a higher level, averaging 8.0 adult female sea lice per fish, FHI requires the creation and implementation of an explicit action plan, agreed with the FHI to effectively reduce and maintain the average number of adult female lice below the reporting level of 3.0. These measures can include the requirement to harvest out the affected fish stocks. 1.3 Current Compliance The recent investments and advancements in better methods and strategies for controlling sea lice have resulted in a trend of progressive improvement in compliance rates with Marine Scotland sea lice reporting and intervention thresholds ( Figure 1). Figure 1. Adult Female Lice Levels vs Marine Scotland Thresholds 2016-2018. 2 Historic Sea Lice Challenges The years 2013-2015 and the beginning of 2016 were challenging for the aquaculture sector as a whole. Elevated summer temperatures accelerated the life cycle of gill parasites and sea lice resulting in higher levels of parasitic loading to the farmed stock (Murray, 2016). This was
5 | 16 during a period when there was a significantly lower cleaner fish capacity and when non- medicinal treatment systems were still being developed. Figure 2. Average monthly adult female levels 2014-2018. 2.1 Marine Harvest Response to Rising Sea Lice Numbers A successful lice management strategy requires access to a range of effective, unrelated active ingredients and tools, to maintain efficacy. There are presently five active ingredient products available in various product formulations for use as sea lice medicines in Scotland; the bath treatments: cypermethrin, azamethiphos, deltamethrin, and hydrogen peroxide; and the in-feed treatment emamectin benzoate. The company invested heavily in improving and developing new tools for better lice management, resulting in a corresponding improvement in lice numbers. This included the increased deployment of both existing, and proven preventative measures. These included non-medical treatments options such as: The first production versions of the new ‘flusher’ systems (Hydrolicer, Thermolicer); The newly developed use of freshwater treatments; and Investment to increase the capacity of biological controllers, such as cleaner fish Ballan Wrasse, Lumpsuckers).
6 | 16 3 Development of Lice Management Tools 3.1 Cleaner Fish The term ‘cleaner fish’ refers to fish, typically Ballan Wrasse and Lumpsuckers, that provide a symbiotic ‘ cleaner’ service to other fish species by the removal of parasites ( Imsland et al., 2014). The Scottish Salmon industry is increasingly stocking cleanerfish into marine salmon farms and they have proven effective in reducing sea lice numbers, and significantly reducing the need for medicinal treatments ( Deady et al., 1995; Leclercq et al., 2014). Currently 100% of all Lumpsuckers deployed per annum in Marine Harvest sites are of farmed origin; however, at present only 10% of the Ballan Wrasse used are of farmed origin. In response to the limited availability of farmed stocks of Ballan Wrasse, Marine Harvest has also now committed to the development of in-house cleaner fish farming. The company has invested significant financial and technical resources into the development of several pioneering cleaner fish hatcheries at locations around the UK. Recently a new recirculation plant was purchased to produce farmed Ballan Wrasse. This facility will be developed over the next few years into a state of the art cleaner fish hatchery. The facility will aim to produce Ballan Wrasse as a sustainable cleaner fish that will then be deployed into the company’s salmon farms. When fully operational, by 2021, this facility will produce 100% of Ballan Wrasse demand for Marine Harvest. In the intervening period there will be a steady transition towards producing cleaner fish stocking requirements from hatchery sources with an annual corresponding declining impact on sourcing these fish from the wild stocks. The sector as a whole is in the process of developing knowledge of growing cleaner fish in hatchery environments. The husbandry methods used to grow, handle and distribute cleaner fish are evolving with the increased knowledge and expertise within this new area. In the interim period, company policy states that wild juvenile Ballan Wrasse less than 12cm in length or mature Ballan Wrasse greater than 22cm in length will not be removed from the wild to ensure the sustainable continuation of viable breeding wild populations of the species. Figure 3. Marine Harvest cleaner fish deployment plan 2018-2022.
7 | 16 3.2 Non-Medicinal Lice Treatments Since late 2015, Marine Harvest Scotland has been developing and investing heavily in the development of non-medicinal methods of sea lice control. Non-medicinal treatments developed and adopted since 2015 include hydrolicer systems; thermolicer system and freshwater treatments. These technologies are still emerging and are constantly being developed, however to date they have demonstrated high treatment efficacies against lice infection. 3.2.1 Hydrolicer Units Hydrolicer systems work by using variable-pressure seawater to dislodge sea lice from the salmon without any detrimental impact on the fish. The method is benign in that it only uses seawater and fish are not exposed to physiological stress or deteriorating water quality. Dislodged sea lice are removed from the used seawater by fine mesh filtration, collected and disposed of by incineration on land. The first prototype hydrolicer unit in Marine Harvest was brought into operation in late December 2015. Given the success of the hydrolicer method, Marine Harvest has invested in developing and expanding the number of hydrolicer units that it now operates. At present, there are three operational hydrolicer units, fitted to the decks of workboats, enabling scheduled and responsive treatments to be undertaken at seawater farm locations. Further development in vessel technology will see hydrolicer units incorporated into the design of well boats or larger dedicated service vessels, further optimising system performance. Marine Harvest currently has the capacity to treat approximately 31,000t in 14 days with mechanical tools i.e. hydrolicers and thermolicers. This equates to 2300t per day. This capacity means the company’s complete standing biomass currently held in the sea can be treated in less than 14 days with non-medicinal treatments. 3.2.2 Thermolicer Units Thermolicers are an effective and safe option for the mechanical removal of sea lice. Sea lice have a low tolerance to changes in body temperature ( Costello, 2006; Torrissen et al., 2013), the system works by exposing the fish to lukewarm water for 30 seconds, which subsequently dislodges sea lice due to the rapid temperature change. Due to the significant body mass of the salmon, relative to the diminutive size of the sea lice, there is no measurable change to body temperature of salmon but sea lice are sufficiently affected to detach from the salmon. Figure 4 illustrates how the system operates.
8 | 16 Figure 4. Schematic of a thermolicer system used within Marine Harvest. The first thermolicer was brought into operation in 2016. Efficacy on all mobile stages of sea lice has generally been very good and fish rarely show negative signs that can be ascribed to the system itself. Recognising the success of this method, a second thermolicer unit was purchased and brought into action in November 2017. The efficacy of the hydro- and thermolicer systems to date are summarised in Figure 5.
9 | 16 Figure 5. Flusher results from 2016 to 2018 for adult female lice clearance levels. 3.2.3 Freshwater Treatments Freshwater treatments have been recognised as an effective strategy for both sea lice and Amoebic Gill Disease (AGD) control. In combination with other available technologies, such as mechanical and biological control, freshwater treatments expand the available fish welfare management tools for sea lice infestation and AGD. Wellboats have been targeted as the optimum solution to deliver freshwater treatment solutions to site, as this offers an opportunity to control conditions, facilitate fish movements, and prevent any mixing of freshwater and seawater during treatment. The majority of treatments are carried out with a holding time of twelve to sixteen hours with lice clearance efficacy regularly high for all stages of sea lice development. Sourcing freshwater abstraction sites for direct fill to well boats is an important element in ensuring the success of this strategy. Currently, the most effective option for freshwater supply is via abstraction from natural freshwater bodies ( burns or lochs). All abstractions require a license from SEPA and the relevant planning authority; to ensure that the rate of abstraction from individual waterbodies is controlled and to ensure good environmental flow standards in the watercourse, maintaining the ecological quality of the watercourse. The abstracted water is then pumped or gravity fed by pipe for containment in enclosed tarpaulins within pens held in seawater, usually located near the abstraction source. In addition to freshwater abstraction, Marine Harvest has invested in a new wellboat, specifically designed to undertake freshwater treatments at seawater farms using the process of desalination. This boat can also, if required, obtain water from a natural freshwater abstraction sources as highlighted above. At present, Marine Harvest have brought into operation five active freshwater abstraction locations and storage pen infrastructure, enabling direct fill of freshwater to wellboats to facilitate sea lice and AGD treatments. There is currently a fleet of three wellboats in Scotland
10 | 16 capable of carrying out freshwater treatments. Several other freshwater abstraction locations in strategic locations to Marine Harvest farms have been identified and relevant permissions are in the process of being secured. 3.2.4 Pen Modifications Research has shown a negative correlation between the swimming depth of fish, and lice prevalence and number within the water column ( Bui et al., 2016; Frenzl et al., 2014). Accordingly, a “deep-strategy” concept has been under development with the aim of keeping fish away from the higher densities of the infective stages of sea lice in the surface layer for most parts of the day, while protecting fish when they need to ascend, e.g. to fill the swim bladder. Such a strategy has the potential to significantly reduce the need for lice treatments, thereby reducing the risk of resistance and improving general fish welfare. There are various aspects to this strategy and different combinations are in place across existing farms, with further research ongoing to optimising effectiveness. Sea Lice Skirts The highest concentrations of sea lice are typically found in the upper parts of the water column and preventing this water from passing through the salmon pens has been found to reduce salmon lice infestation ( Frenzl et al., 2014; Hevrøy et al., 2003). A lice skirt is a 6m deep tarpaulin which is placed into the water around a fish pen to a depth of 5m sub surface; 1m of the tarpaulin is retained above the surface of the sea. Lice skirts have been widely implemented by Marine Harvest since 2016. The deployment of tarpaulin skirts has been shown to delay the start of the next “first pen” infection. quality within pens utilising lice skirts is a subject area of ongoing research by Marine Harvest. Deep Feeding and Deep Lights The use of deep feeding equipment that has high capacity and spreads the feed spatially at depth, promoting deep feeding behaviours by salmon and its efficacy as a lice prevention measure is presently the subject of developing research internally. Prior research has also demonstrated that the behaviour of salmon, including swimming depth in sea pens, shows seasonal variations dependent upon a range of environmental factors including light conditions Bui et al., 2016; Frenzl et al., 2014; Hevrøy et al., 2003). During periods of high light intensity, salmon typically swim deeper than in periods of low light intensity where the salmon tend to swim shallower. Marine Harvest are therefore undertaking further research trials on the potential for behavioural manipulation of salmon, and the subsequent lice prevalence and the efficacy against infection by utilising measures such as deep feeding and deep lights. 3.3 Medicinal Treatments The industry adopts a range of measures to control fish health, including good husbandry and management, natural feed additives, site fallowing, mechanical tools and biological tools such as cleaner fish and, where necessary, veterinary medicines. There are presently five active ingredients available (in various product formulations) for use as sea lice medicines in Scotland: the bath treatments: cypermethin, azamethiphos,
11 | 16 deltamethrin, and hydrogen peroxide; and the in-feed treatment emamectin benzoate. All medicines are prescribed by the company veterinarian and their use is regulated by the Veterinary Medicine Directorate ( VMD) as well as SEPA. There are also strict criteria and procedures for monitoring medicinal residues in farmed salmon under food safety regulations as is the case with terrestrial farmed animals. 3.4 Good Husbandry Practice Marine Harvest Scotland’ s goal is to rear healthy fish and maximise the welfare on fish stocks. Fish are tended to under conditions that satisfy their biological needs for food, clean water and space, and it is ensured that the fish obtain the necessary nutrients for good health throughout production. The fish are stocked at densities that balance welfare and enhance performance. Coordinated fallowing and synchronised production are integral components of Marine Harvest’s farming practices, which reduce biological risk. Under the supervision of fish health professionals, Marine Harvest continuously apply good farming practices and high standards of biosecurity to optimize the health and welfare of stocks. By adhering to stringent veterinary health plans and recognized fish welfare standards, such as Royal Society for the Prevention of Cruelty to Animals ( RSPCA), and vaccinating 100% of our fish, we reduce biological and health risks. Interventions, such as biological control, medicinal treatments, and thermic/ physical removal of sea lice are only part of an integrated control strategy and would not be successful without a firm foundation of good fish welfare, husbandry and other preventative measures which include fallow periods, single year class sites, regular lice counting, etc. Fallow Periods The aim is to deprive sea lice of available hosts through fallow periods, i.e. no farmed salmonids in a farm area for at least four weeks, is highly effective in re-setting the level of sea lice, as free-swimming juvenile infective stages will die within one week if they can’t find hosts Torrissen et al., 2013). Current policy states that each site is fallowed for at least six weeks between production cycles. Single year class sites This is linked to fallow period and requires all sites to be stocked with new smolts within a defined period which allows all to be harvested out again in time for the agreed fallow. New smolts should not be introduced to a site that still holds harvest sized fish from the previous generation. Regular lice counting on farms The Code of Good Practice requires farms to collect representative lice data at least once a week from all stocked farms to monitor and actively manage sea lice levels in their stock CoGP, 2003). Marine Harvest views this as critical to success and dedicates extensive resource to gathering this data at an even higher level of resolution (more fish counted, from more pens) than required by the COGP.
12 | 16 Communication between companies Marine Harvest strongly believes that communication with the Health Teams of neighbouring farms and companies is important, and has a policy of non competative business transparency in this regard. 4 Treatment Selection and Hierarchy The criteria defining treatments selected for a particular site is determined on the basis of a number of decision criteria. Physical systems, i.e. lice treatment skirts are typically installed throughout the production cycle. Specific strategic treatments are applied to achieve defined levels of protection, for example the in-feed treatment emamectin benzoate is frequently applied at the beginning of the wild smolt migration season to maximum protection against infection during this sensitive period. Decisions regarding the use of other measures are based on a series of principals to ensure a diverse range or treatments are applied to ensure continued treatment efficacy, the type of treatment last applied and level of efficacy achieved, and the availability and capacity of specific treatment options etc. Given the level of responsiveness required by a lice treatment plan, a defined treatment strategy is unlikely to be as effective and could lead to the inefficient or unnecessary use of management tools. Therefore, an example of a treatment plan for the proposed site is provided in Appendix 1 of this document. 5 Conclusion Effective sea lice management is important for the health and welfare of farmed fish, and also to reduce the risk of fish farms on wild salmonids. The company has invested heavily, and continues to invest, in the management and advancement of tools and methodologies to combat the occurrence of sea lice on its farms. Measures implemented include cleaner fish, hydrolicers, thermolicers and freshwater treatments. These new measures are in parallel with new internal policies An example treatment plan for the site is provided illustrating the typical timing, nature and range of treatments implemented during a production cycle to demonstrate the extent of the potential tools available to ensure sea lice levels meet internal and external targets.
13 | 16 Appendix 1 Example Treatment Plan: Loch Alsh Ten 120m Circle Pens 2500 tonne Maximum Standing Biomass Note: The company promotes proactive sealice control, as well as monitoring and intervention as part of a reactive approach. This document is purely an example of how lice levels could be managed based on the current company budget, experience, and guidance such as the National Treatment Strategy. The need for intervention, the method used, and the timing is dependent on site conditions and thus subject to change following the results of regular monitoring. Further resource could be made available in response to fish health. Available Treatments Colour Code Lice Skirts In-Feed Bath Cleaner Fish Mechanical Production Year Calander Cycle Treatment Quarter Week Week Type 1 1 Lice Skirts/ Cleaner Fish 2 2 3 3 4 4 5 5 6 6 19 Q1 7 7 8 8 9 9 10 10 11 11 12 12 13 13 14 14 15 15 16 16 17 17 18 18 19 19 19 Q2 20 20 21 21 In-feed 22 22 23 23 Cleanerfish 24 24 25 25 26 26 27 27 Bath 28 28 19 Q3 29 29 30 30 31 31
14 | 16 32 32 33 33 34 34 35 35 36 36 Mechanical 37 37 38 33 39 34 40 35 41 36 42 37 43 38 44 39 45 40 19 Q4 46 41 47 42 48 43 49 44 50 45 51 46 52 47 53 48 54 49 55 50 56 51 57 52 58 53 20 Q1 59 54 60 55 61 56 62 57 63 58 64 59 65 60 Bath 66 61 67 62 68 63 69 64 70 65 71 66 20 Q2 72 67 73 68 Mechanical 74 69 75 70 76 71 77 72 78 73 79 74 80 75 81 76 82 77 Mechanical 20 Q3 83 78 84 79 85 80 86 81 87 82
15 | 16 88 83 Mechanical 89 84 90 85 91 86 92 87 93 88 94 89 95 90 96 91 97 92 20 Q4 98 93 99 94 100 95 101 96 102 97 103 98 104 99
16 | 16 6 References Bui, S., Oppedal, F., Stien, L., Dempster, T., 2016. Sea lice infestation level alters salmon swimming depth in sea-pens. Aquac. Environ. Interact. 8, 429–435. doi:10.3354/aei00188 CoGP, 2003. Code of Good Practice: Chapter 4 Seawater Lochs. http://thecodeofgoodpractice. co.uk/chapters/ chapter- 4-seawater- lochs/ 2003. Costello, M.J., 2006. Ecology of sea lice parasitic on farmed and wild fish. Trends Parasitol. 22, 475–483. doi:10.1016/j.pt.2006.08.006 Deady, S., Varian, S.J.A., Fives, J.M., 1995. The use of cleaner- fish to control sea lice on two Irish salmon (Salmo salar) farms with particular reference to wrasse behaviour in salmon pens. Aquaculture 131, 73–90. doi:10.1016/0044-8486(94)00331-H Frenzl, B., Stien, L.H., Cockerill, D., Oppedal, F., Richards, R.H., Shinn, A.P., Bron, J.E., Migaud, H., 2014. Manipulation of farmed Atlantic salmon swimming behaviour through the adjustment of lighting and feeding regimes as a tool for salmon lice control. Aquaculture 424–425, 183–188. doi:10.1016/j.aquaculture.2013.12.012 Hevrøy, E.M., Boxaspen, K., Oppedal, F., Taranger, G.L., Holm, J.C., 2003. The effect of artificial light treatment and depth on the infestation of the sea louse Lepeophtheirus salmonis on Atlantic salmon ( Salmo salar L.) culture. Aquaculture 220, 1–14. doi:10.1016/S0044-8486(02)00189-8 Imsland, A.K., Reynolds, P., Eliassen, G., Hangstad, T.A., Foss, A., Vikingstad, E., Elvegård, T.A., 2014. The use of lumpfish ( Cyclopterus lumpus L.) to control sea lice Lepeophtheirus salmonis Krøyer) infestations in intensively farmed Atlantic salmon Salmo salar L.). Aquaculture 424–425, 18–23. doi:10.1016/j.aquaculture. 2013.12.033 Leclercq, E., Davie, A., Migaud, H., 2014. Delousing efficiency of farmed ballan wrasse (Labrus bergylta) against Lepeophtheirus salmonis infecting Atlantic salmon (Salmo salar) post- smolts. Pest Manag. Sci. 70, 1274–1282. doi:10.1002/ps.3692 Murray, A.G., 2016. A note on sea lice abundance on farmed Atlantic salmon in Scotland 2011- 2013: Significant regional and seasonal variation. Aquac. Res. 47, 961–968. doi:10.1111/are.12554 Powell, M.D., Reynolds, P., Kristensen, T., 2015. Freshwater treatment of amoebic gill disease and sea-lice in seawater salmon production: Considerations of water chemistry and fish welfare in Norway. Aquaculture 448, 18–28. doi:10.1016/j.aquaculture. 2015.05.027 Torrissen, O., Jones, S., Asche, F., Guttormsen, A., Skilbrei, O.T., Nilsen, F., Horsberg, T.E., Jackson, D., 2013. Salmon lice - impact on wild salmonids and salmon aquaculture. J. Fish Dis. 36, 171–194. doi:10.1111/jfd.12061
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