Scottish Animal Welfare Commission: welfare of cleaner fish used in the Scottish salmon industry - report

7. References

Barrett, L.K., Oppedal, F., Robinson, N., Dempster, T. (2020a). Prevention not cure: a review of methods to prevent sea lice infestation in salmon aquaculture. Reviews in Aquaculture 12(4): 2547-2543.

Barrett, L.T., Overton, K., Stien, L.H., Oppedal, F., Dempster, T. (2020b). Effect of cleaner fish on sea lice in Norwegian salmon aquaculture: a national scale data analysis. International Journal for Parasitology 50:787-796.

Boerlage, A.S., Shrestha, S., Leinonen, I., Jansen, M.D., Revie, C.W., Reeves, A., Toma, L. (2024). Sea lice management measures for farmed Atlantic salmon (Salmo salar) in Scotland: Costs and effectiveness. Aquaculture 580 (1), 740274.

Bowers JM, Mustafa A, Speare DJ, Conboy GA, Brimacombe M, Sims DE, Burka JF (2000) The physiological response of Atlantic salmon, Salmo salar L., to a single

experimental challenge with sea lice, Lepeophtheirus salmonis. J Fish Dis 23: 165−172

Brooker, AJ., Papadopoulou, A., Gutierrez, C., Rey, S., Davie, A., Migaud, H.

(2018) Sustainable production and use of cleaner fish for the biological control of sea lice: recent advances and current challenges. Veterinary Record 183, 383

Brooker, A.J., Davie, A., Leclercq, E., Zerafa, B and Migaud, H. (2020). Pre-deployment acclimatisation of farmed ballan wrasse (Labrus bergylta) to sea-cage conditions promotes behaviour analogous to wild conspecifics when used as cleaner fish in Atlantic salmon (Salmo salar) farms. Aquaculture 520, 734771

Brooker, A., Jimenez, E., Bennett, L. (2021). Aquawrasse: improving delousing behaviour and efficacy in farmed ballan wrasse. Report to SAIC.

Cai, W., Zhong, L., Parrish, K., Kumar, S., Eslamloo, K., Fajei, E., Whyte, S.K., Purcell, S.L., Jahangiri, L., Li, R., Caballero Solares, A., Taylor, R.G., Balder, R., Rise, M.L., Fast, M.D. (2024). Synergies of co-infecting pathogens, sea lice (Lepeophetheirus salmonis) and Moritella viscosa, are impacted by exposure order, and host response to initial infection. Aquaculture 591, 741115

Dahle, O.M.V., Blindheim, S.H., Nylund, A., Karlsbakk, E., Breck, O., Glosvik, H., Andersen, L. (2020). Atlantic salmon Salmo salar and ballan wrasse Labrus bergylta display different susceptibility to clonal strains of Paramoeba perurans. Diseases of Aquatic Organisms 140:55-72.

Geitung, L., Wright, D.W., Oppedal, F., Stien, L.H., Vågseth, T., Madaro, A., (2020). Cleaner fish growth, welfare and survival in Atlantic salmon sea cages during an autumn-winter production. Aquaculture 528, 735623

Grant, B., Davie, A., Taggart, J.B., Selly, S.L., Picchi, N., Bradley, C., Prodohl, P., Leclercq, E. and Migaud, H. (2016). Seasonal changes in broodstock spawning performance and egg quality in ballan wrasse (Labrus bergylta). Aquaculture 464: 505-514.

Grimnes A, Jakobsen PJ (1996) The physiological effects of salmon lice infection on post-smolt of Atlantic salmon. Journal of Fish Biology 48: 1179–1194

Hall, L.M., Smith, R.J., Munro, E.S., Matejusova, I., Allan, C.E.T., Murray, A.G., Duguid, S.J., Salama, N.K.G., McBeath, A.J.A., Wallace, I.S., Bain,N., Marcos-Lopez, M. and Raynard, R.S. (2013). Epidemiology and control of an outbreak of viral haemorrhagic septicaemia in wrasse around Shetland commencing 2012. Scottish Marine and Freshwater Science 4 No. 03. Published by Marine Scotland – Science ISSN: 2043-7722

Helland, S., Dahle, S. V.W., Hough, C.,Borthen, J., (2014). Production of ballan wrasse. Science and practice. Fiskeri- og havbruksnæringens forskningsfond (FHF) - The Norwegian Seafood Research Fund.

Imsland, A.K., Reynolds, P., Eliassen, G., Mortensen, A., Hansen, O., Puvanedndran, V., Hangstand, T.A., Jónsdóttir, Ó.D.B., Emaus, P-A., Elvegård, T.A., Lemmens, S.C.A., Rydland, R., Nytro., A.V., and Jonassen, T.M. (2016). Is cleaning behaviour in lumpfish (Cyclopterus lumpus) parentally controlled? Aquaculture 459 156-165.

Jimenez, D.F., Revie, C.W., Hardy, S.P., Jansen P.A., and Gettinby, G. (2013) Multivariate evaluation of the effectiveness of treatment efficacy of cypermethrin against sea lice (Lepeophtheirus salmonis) in Atlantic salmon (Salmo salar). BMC Vet Res 9, 258.

Leclerq, E., Grant, B., Davie, A., Migaud, H. (2014). Gender distribution, sexual size dimorphism and morphometric sexing in ballan wrasse Labrus bergylta. Journal of Fish Biology, 84: 1842-1862.

Leclercq, E., Zerafa, B., Brooker, A.J., Davie, A., and Migaud, H. (2018). Application of passive-acoustic telemetry to explore the behaviour of ballan wrasse (Labrus bergylta) and lumpfish (Cyclopterus lumpus) in commercial Scottish salmon sea-pens. Aquaculture 495: 1-12.

Leggatt, R.A., Scheer, K.W., Afonso, L.O.B. and Iwana, G.K. (2006). Triploid and diploid rainbow trout do not differ in their stress response to transportation. North American Journal of Aquaculture, 68: 1-8.

Lie, K.K., Tørresen, O.K., Solbakken, M.H., Rønnestad, I., Tooming-Klunderud, A., Nederbragt, A.J., Jentoft, S. and Sæle, Ø. (2018). Loss of stomach, loss of appetite? Sequencing of the ballan wrasse (Labrus bergylta) genome and intestinal transcriptomic profiling illuminate the evolution of loss of stomach function in fish. BMC Genomics, 19: 1-17.

Marine Directorate, Scottish Government, (2022b). Fishing – wild wrasse: application for a letter of derogation.

Muncaster, S., Norberg, B. and Andersson, E., (2013). Natural sex change in the temperate protogynous Ballan wrasse Labrus bergylta. Journal of Fish Biology 82: 1858-1870.

Nilsen A, Viljugrein H, Røsæg M, Colquhoun D (2014). Rensefiskhelse—kartlegging av dødelighet og dødelighetsårsaker.Veterinærinstituttets rapportserie 12-2014. Veterinærinstituttet, Oslo

Nilsson, J., Moltumyr, L., Madaro, A., Kristiansen, T.S., Gåsnes, S.K., Mejdell, C.M., Gismervik, K., Helge Stien, J.H. (2019). Sudden exposure to warm water causes instant behavioural responses indicative of nociception or pain in Atlantic salmon. Veterinary and Animal Science. vol 8, 100076.

Noble, C., Gismervik, K., Iversen, M. H., Kolarevic, J., Nilsson, J., Stien, L. H. &

Turnbull, J. F. (Eds.) (2018). Welfare Indicators for farmed Atlantic salmon: tools for assessing fish welfare 351pp.

Noble, C., Iversen, M. H., Lein, I., Kolarevic, J., Johansen, L. –H., Burgerhout, E., Puvanendran, V., Kousoulaki, K., Aas, G. H., Stene, A. & Espmark, Å. M. (2019). RENSVEL OWI FACT SHEET SERIES: An introduction to Operational and Laboratory based Welfare Indicators for ballan wrasse (Labrus bergylta). 43 pp.

Norland, S., Saele O., Ronnestand, I., (2022). Developmental stages of the ballan wrasse from first feeding through metamorphosis: Cranial ossification and the digestive system. Journal of Anatomy 241: 337–357.

Overton, K., Dempster, T., Oppedal, F., Kristiansen, T.S., Gismervik, K., Stien, L.H. (2019). Salmon lice treatments and salmon mortality in Norwegian aquaculture: a review. Reviews in Aquaculture 11(4): 1398-1417.

Overton, K., Barrett, L.T., Oppedal, F., Kristiansen, T.S., Dempster, T. (2020). Sea lice removal by cleaner fish in salmon aquaculture: a review of the evidence base. Aquaculture Environment Interactions 12: 31–44.

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.

Powell, A., Treasurer, J.W., Pooley, C.L., Keay, A.J., Lloyd, R., Imsland, A.K., De Leaniz, C.G. (2017). Use of lumpfish for sea-lice control in salmon farming.

Pritchard, C.J., Salama, N.K.G., Berrill, I., Martin, S.A.M., Marshall, C.T. (2025a). Investigating the relative vulnerability of a temperate coastal fish community to a live wrasse pot fishery. Fisheries Research 284: 107327.

Pritchard, C.J., Treasurer, J., Highway, A., Salama, N.K.G., Berrill, I., Martin, S.A., Marshall, C.T. (2025b). Preliminary investigation into the appropriateness of control measures and harvest control rules for a data-limited, sex-changing fish managed by harvest slot limits. Aquaculture, Fish and Fisheries. 5:e70091.

RSPCA, 2024. RSPCA welfare standards: Farmed Atlantic salmon. Horsham RH12 1GY, United Kingdom.

Scottish Government (2022). Fishing – wild wrasse: application for a letter of derogation.

Scottish Government (2024a). Scottish Fish Farm Production Survey 2023. St Andrews House, Edinburgh, EH1 3DG. ISBN: 978-1-83601-921-3 (web only).

Scottish Government (2024b). Wild wrasse fishery – 2021/22 Report. St Andrews House, Edinburgh, EH1 3DG

Scottish Government (2025), Regulated commercial use of traps/pots to catch wrasse in Scottish inshore waters. Fisheries Assessment. St Andrews House, Edinburgh, EH1 3DG. ISBN: 978-1-83691-776-2 (web only).

Scottish Finfish Aquaculture (2016). Code of Good Practice.

Skiftesvik, A.B., Bjelland, R.M., Durif, C., Johansen, I.S., Browman, H. (2013). Delousing of Atlantic salmon (Salmo salar) by cultured vs. wild ballan wrasse (Labrus bergylta). Aquaculture 402-403: 113-118.

Soares, M.C., Bshary, R., Mendonça, R., Grutter, A.S., Oliveira, R.F. (2012) Arginine vasotocin regulation of interspecific cooperative behaviour in a cleaner fish. PLoS ONE 7(7): e39583.

Treasurer, J. W. (2002) A review of potential pathogens of sea lice and the application of cleaner fish in biological control. Pest Management Science 58, 546–558

Treasurer, J. (2018). An introduction to sea lice and the rise of cleaner fish. In: Cleaner Fish Biology and Aquaculture Applications. Ed: J. Treasurer. 5M Publishing Ltd. Sheffield, England

Treasurer, J., Prickett, R., Zietz, M., Hempleman, C., Carcia de Leaniz, C. (2024). Cleaner fish rearing and deployment in the UK. Chapter 19 In: Cleaner fish biology and aquaculture applications. CABI. Ed: J. Treasurer

Truskanov, N., Emery, Y., and Bshary, R. (2020). Juvenile cleaner fish can socially learn the consequences of cheating. Nature Communications 11:1159