Migration routes and behaviour of Atlantic salmon and sea trout around operational wind turbine arrays in the Moray Firth, Scotland

References

Akaike, H. (1973). Information theory and an extension of the maximum likelihood principle. In B. N. Petrov & F. Csaki (Eds.), Second International Symposium on Information Theory (Vol. 1, pp. 267–281). Springer Verlag.

Arnett, E. B., & May, R. F. (2016). Mitigating wind energy impacts on wildlife: Approaches for multiple taxa. Human-Wildlife Interactions, 10(1).

Artero, C., Marsh, J. E., Beaumont, W. A., Lecointre, T., & Lauridsen, R. B. (2025). Spatial–temporal variation in the vertical behaviour of sea trout kelts during their marine phase. Reviews in Fish Biology and Fisheries, 35(3), 1407–1427.

Barton, B., De Dominicis, M., O’Hara Murray, R., & Campbell, L. (2022). Scottish shelf model 3.02--27 Year reanalysis. National Oceanography Centre, Liverpool, United Kingdom.

Bicknell, A. W. J., Gierhart, S., Newton, M., Main, R., Thompson, P., & Witt, M. J. (2025). The role of acoustic telemetry to assess the effects of offshore wind infrastructure on fish behaviour, populations and predation. Renewable and Sustainable Energy Reviews, 212, 115306.

Bicknell, A. W. J., Gierhart, S., & Witt, M. J. (2025). Site and species dependent effects of offshore wind farms on fish populations. Marine Environmental Research, 205, 106977.

Birnie-Gauvin, K., Thorstad, E. B., & Aarestrup, K. (2019). Overlooked aspects of the Salmo salar and Salmo trutta lifecycles. Reviews in Fish Biology and Fisheries, 29(4), 749–766.

Brownscombe, J. W., Griffin, L. P., Chapman, J. M., Morley, D., Acosta, A., Crossin, G. T., Iverson, S. J., Adams, A. J., Cooke, S. J., & Danylchuk, A. J. (2019). A practical method to account for variation in detection range in acoustic telemetry arrays to accurately quantify the spatial ecology of aquatic animals. Methods in Ecology and Evolution, 0(ja).

Brunnschweiler, J. M. (2009). Tracking free-ranging sharks with hand-fed intra-gastric acoustic transmitters. Marine and Freshwater Behaviour and Physiology, 42(3).

Butler, J. R. A., & Walker, A. F. (2007). Characteristics of the Sea Trout Salmo trutta (L.) Stock Collapse in the River Ewe (Wester Ross, Scotland), in 1988-2001. In Sea Trout: Biology, Conservation and Management (pp. 45–59).

Coolen, J. W. P., Van Der Weide, B., Cuperus, J., Blomberg, M., Van Moorsel, G. W. N. M., Faasse, M. A., Bos, O. G., Degraer, S., & Lindeboom, H. J. (2020). Benthic biodiversity on old platforms, young wind farms, and rocky reefs. Journal of Marine Science, 77(3).

Coolen, J. W. P., Vanaverbeke, J., Dannheim, J., Garcia, C., Birchenough, S. N. R., Krone, R., & Beermann, J. (2022). Generalized changes of benthic communities after construction of wind farms in the southern North Sea. Journal of Environmental Management, 315.

Dadswell, M., Spares, A., Reader, J., McLean, M., McDermott, T., Samways, K., & Lilly, J. (2022). The Decline and Impending Collapse of the Atlantic Salmon (Salmo salar) Population in the North Atlantic Ocean: A Review of Possible Causes. In Reviews in Fisheries Science and Aquaculture (Vol. 30, Issue 2).

Dannheim, J., Bergström, L., Birchenough, S. N. R., Brzana, R., Boon, A. R., Coolen, J. W. P., Dauvin, J. C., De Mesel, I., Derweduwen, J., Gill, A. B., Hutchison, Z. L., Jackson, A. C., Janas, U., Martin, G., Raoux, A., Reubens, J., Rostin, L., Vanaverbeke, J., Wilding, T. A., … Degraer, S. (2020). Benthic effects of offshore renewables: Identification of knowledge gaps and urgently needed research. In ICES Journal of Marine Science (Vol. 77, Issue 3).

Davidsen, J. G., Plantalech Manel-La, N., Økland, F., Diserud, O. H., Thorstad, E. B., Finstad, B., Sivertsgård, R., McKinley, R. S., & Rikardsen, A. H. (2008). Changes in swimming depths of Atlantic salmon Salmo salar post-smolts relative to light intensity. Journal of Fish Biology, 73(4).

Diadromous Fish ScotMER Receptor Group. (2024). Diadromous Fish Evidence Map.

Eldøy, S. H., Davidsen, J. G., Thorstad, E. B., Whoriskey, F., Aarestrup, K., Næsje, T. F., Rønning, L., Sjursen, A. D., Rikardsen, A. H., & Arnekleiv, J. V. (2015). Marine migration and habitat use of anadromous brown trout (Salmo trutta). Canadian Journal of Fisheries and Aquatic Sciences, 72 (9), 1366–1378.

Eldøy, S. H., Davidsen, J. G., Thorstad, E. B., Whoriskey, F. G., Aarestrup, K., Næsje, T. F., Rønning, L., Sjursen, A. D., Rikardsen, A. H., & Arnekleiv, J. V. (2017). Marine depth use of sea trout Salmo trutta in fjord areas of central Norway. Journal of Fish Biology, 91(5).

Gargan, P. G., Poole, W. R., & Forde, G. P. (2007). A Review of the Status of Irish Sea Trout Stocks. In Sea Trout: Biology, Conservation and Management.

Gilbey, J., Utne, K. R., Wennevik, V., Beck, A. C., Kausrud, K., Hindar, K., Garcia de Leaniz, C., Cherbonnel, C., Coughlan, J., Cross, T. F., Dillane, E., Ensing, D., García-Vázquez, E., Hole, L. R., Holm, M., Holst, J. C., Jacobsen, J. A., Jensen, A. J., Karlsson, S., … Verspoor, E. (2021). The early marine distribution of Atlantic salmon in the North-east Atlantic: A genetically informed stock-specific synthesis. Fish and Fisheries, 22(6).

Gill, A. B., Bartlett, M., & Thomsen, F. (2012). Potential interactions between diadromous fishes of U.K. conservation importance and the electromagnetic fields and subsea noise from marine renewable energy developments. Journal of Fish Biology, 81(2).

Gillson, J. P., Bašić, T., Davison, P. I., Riley, W. D., Talks, L., Walker, A. M., & Russell, I. C. (2022). A review of marine stressors impacting Atlantic salmon Salmo salar, with an assessment of the major threats to English stocks. In Reviews in Fish Biology and Fisheries (Vol. 32, Issue 3).

GWEC. (2025). Global Offshore Wind Report 2025. In Global Wind Energy Council.

Hansen, L. P., Jonsson, N., & Jonsson, B. (1993). Oceanic migration in homing Atlantic salmon. Animal Behaviour, 45(5).

Harding, H., Bruintjes, R., Radford, A., & Simpson, S. (2016). Measurement of Hearing in the Atlantic salmon (Salmo salar) using Auditory Evoked Potentials, and effects of Pile Driving Playback on salmon Behaviour and Physiology. Scottish Marine and Freshwater Science Report Vol 7 No 11.

Holm, M., Holst, J. C., & Hansen, L. P. (2000). Spatial and temporal distribution of post-smolts of Atlantic salmon (Salmo salar L.) in the Norwegian Sea and adjacent areas. ICES Journal of Marine Science, 57(4).

Honkanen, H., Moore, I., Roger, J., Adams, C. E., & Dodd, J. (2024). Diadromous Fish in the Context of Offshore Wind - Review of current knowledge & future research.

ICES. (2013). Report of the Workshop on Sea Trout (WKTRUTTA), 12–14 November 2013, ICES Headquarters, Copenhagen, Denmark. ICES CM 2013/SSGEF:15. 243 pp.

International Energy Agency. (2023). Renewables 2022 - Analysis and forecast to 2027.

Iorio-Merlo, V., Fernandez-Betelu, O., Benhemma-Le Gall, A., Graham, I. M., & Thompson, P. M. (2023). Task 4.2. Work Package 4 – Changes in the occurrence of harbour porpoises following the construction of Moray Firth offshore windfarms. PrePARED Report, No. 002. March 2024.

Jacoby, D. M. P., & Piper, A. T. (2025). What acoustic telemetry can and cannot tell us about fish biology. In Journal of Fish Biology (Vol. 106, Issue 5).

Jepsen, N., Aarestrup, K., Økland, F., & Rasmussen, G. (1998). Survival of radio-tagged Atlantic salmon (Salmo salar (L.)) and trout (Salmo trutta L.) smolts passing a reservoir during seaward migration. Hydrobiologia, 371–372, 347–353.

Katzner, T. E., Nelson, D. M., Marques, A. T., Voigt, C. C., Lambertucci, S. A., Rebolo, N., Bernard, E., Diehl, R., & Murgatroyd, M. (2025). Impacts of onshore wind energy production on biodiversity. Nature Reviews Biodiversity, 1(9), 567–580.

Klimley, A. P., Wyman, M. T., & Kavet, R. (2017). Chinook salmon and green sturgeon migrate through San Francisco Estuary despite large distortions in the local magnetic field produced by bridges. PLoS ONE, 12(6).

Klinard, N. V., & Matley, J. K. (2020). Living until proven dead: addressing mortality in acoustic telemetry research. In Reviews in Fish Biology and Fisheries (Vol. 30, Issue 3).

Knutsen, J. A., Knutsen, H., Gjøsæter, J., & Jonsson, B. (2001). Food of anadromous brown trout at sea. Journal of Fish Biology, 59(3).

Kristensen, M. L., Righton, D., Del Villar-Guerra, D., Baktoft, H., & Aarestrup, K. (2018). Temperature and depth preferences of adult sea trout Salmo trutta during the marine migration phase. Marine Ecology Progress Series, 599

Kristensen, M. L., Righton, D., Villar-Guerra, D. del, Baktoft, H., & Aarestrup, K. (2019). Behaviour of adult sea trout Salmo trutta that survive or die at sea. Estuarine, Coastal and Shelf Science, 227.

Lacroix, G. L., McCurdy, P., & Knox, D. (2004). Migration of Atlantic Salmon Postsmolts in Relation to Habitat Use in a Coastal System. Transactions of the American Fisheries Society, 133(6).

Langhamer, O. (2012). Artificial Reef Effect in relation to Offshore Renewable Energy Conversion: State of the Art. The Scientific World Journal, 2012(1), 386713.

Laranjeiro, T., May, R., & Verones, F. (2018). Impacts of onshore wind energy production on birds and bats: recommendations for future life cycle impact assessment developments. In International Journal of Life Cycle Assessment (Vol. 23, Issue 10).

Lennox, R. J., Dahlmo, L. S., Ford, A. T., Sortland, L. K., Vogel, E. F., & Vollset, K. W. (2023). Predation research with electronic tagging. Wildlife Biology, 2023(1).

Lilly, J., Honkanen, H. H., Rodger, J. R., del Villar, D., Boylan, P., Green, A., Pereiro, D., Wilkie, L., Kennedy, R., Barkley, A., Rosell, R., Maoiléidigh, N., O’Neill, R., Waters, C., Cotter, D., Bailey, D., Roche, W., McGill, R., Barry, J., … Adams, C. E. (2024). Migration patterns and navigation cues of Atlantic salmon post-smolts migrating from 12 rivers through the coastal zones around the Irish Sea. Journal of Fish Biology, 104(1).

Lothian, A. J., Newton, M., Barry, J., Walters, M., Miller, R. C., & Adams, C. E. (2018). Migration pathways, speed and mortality of Atlantic salmon (Salmo salar) smolts in a Scottish river and the near-shore coastal marine environment. Ecology of Freshwater Fish, 27(2).

Main, R. A. (2021). Migration of Atlantic salmon (Salmo salar) smolts and post-smolts from a Scottish east coast river [MSc]. University of Glasgow.

Main, R., Reeve, A., Archer, J., O’Hara Murray, R., Newton, M., Gardiner, R., Buddendorf, B., Armstrong, J., Davies, I., & Hawkins, L. (2023). North East Scotland Salmon and Sea Trout Tracking Array. River Dee Trust (RDT) and Marine Scotland Science (MSS).

Malcolm, I., Millar, C., & Millidine, K. (2015). Spatio-temporal variability in Scottish smolt emigration times and sizes. Marine Scotland Science. 10.7489/1590-1.

McLean, D. L., Taylor, M. D., Partridge, J. C., Gibbons, B., Langlois, T. J., Malseed, B. E., Smith, L. D., & Bond, T. (2018). Fish and habitats on wellhead infrastructure on the north west shelf of Western Australia. Continental Shelf Research, 164.

Middlemas, S. J., Armstrong, J. D., & Thompson, P. M. (2007). The Significance of Marine Mammal Predation on Salmon and Sea Trout. In Salmon at the Edge (pp. 41–60).

Nevoux, M., Finstad, B., Davidsen, J. G., Finlay, R., Josset, Q., Poole, R., Höjesjö, J., Aarestrup, K., Persson, L., Tolvanen, O., & Jonsson, B. (2019). Environmental influences on life history strategies in partially anadromous brown trout (Salmo trutta, Salmonidae). Fish and Fisheries, 20(6), 1051–1082.

Newton, M., Barry, J., Lothian, A., Main, R., Honkanen, H., McKelvey, S., Thompson, P., Davies, I., Brockie, N., Stephen, A., Murray, R. O. H., Gardiner, R., Campbell, L., Stainer, P., & Adams, C. (2021). Counterintuitive active directional swimming behaviour by Atlantic salmon during seaward migration in the coastal zone. ICES Journal of Marine Science, 78(5).

Nunn, A. D., Ainsworth, R. F., Walton, S., Bean, C. W., Hatton-Ellis, T. W., Brown, A., Evans, R., Atterborne, A., Ottewell, D., & Noble, R. A. A. (2023). Extinction risks and threats facing the freshwater fishes of Britain. Aquatic Conservation: Marine and Freshwater Ecosystems, 33(12).

Ounsley, J. P., Gallego, A., Morris, D. J., & Armstrong, J. D. (2020). Regional variation in directed swimming by Atlantic salmon smolts leaving Scottish waters for their oceanic feeding grounds - A modelling study. ICES Journal of Marine Science, 77(1).

Pratten, D. J., & Shearer, W. M. (1983). Sea Trout of the North Esk. Aquaculture Research, 14(2).

R Core Team. (2024). R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing.

Rikardsen, A. H., Amundsen, P. A., Knudsen, R., & Sandring, S. (2006). Seasonal marine feeding and body condition of sea trout (Salmo trutta) at its northern distribution. ICES Journal of Marine Science, 63(3).

Rikardsen, A. H., Diserud, O. H., Elliott, J. M., Dempson, J. B., Sturlaugsson, J., & Jensen, A. J. (2007). The marine temperature and depth preferences of Arctic charr (Salvelinus alpinus) and sea trout (Salmo trutta), as recorded by data storage tags. Fisheries Oceanography, 16(5).

Rodger, J. R., Lilly, J., Honkanen, H. M., del Villar, D., Kennedy, R., Maoiléidigh, N. Ó., Boylan, P., Rosell, R., Morris, D. J., O’Neill, R., Waters, C., Cotter, D., Wilkie, L., Barkley, A., Green, A., Beck, S. V, Ribbens, J., Henderson, J., Parke, D., … Adams, C. E. (2025). Inshore and offshore marine migration pathways of Atlantic salmon post-smolts from multiple rivers in Scotland, England, Northern Ireland, and Ireland. Journal of Fish Biology, 106(5), 1422–1439.

Russell, D. J. F., Brasseur, S. M. J. M., Thompson, D., Hastie, G. D., Janik, V. M., Aarts, G., McClintock, B. T., Matthiopoulos, J., Moss, S. E. W., & McConnell, B. (2014). Marine mammals trace anthropogenic structures at sea. In Current Biology (Vol. 24, Issue 14).

Scottish Government. (2023). Draft Energy Strategy and Just Transition Plan.

Scottish Offshore Wind Energy Council (2024) Scotland’s offshore wind industry - Progress and Potential ( 2024 Annual Review).

Skaala, Ø., Kålås, S., & Borgstrøm, R. (2014). Evidence of salmon lice-induced mortality of anadromous brown trout (Salmo trutta) in the Hardangerfjord, Norway. Marine Biology Research, 10(3), 279–288.

Svendsen, J. C., Ibanez-Erquiaga, B., Savina, E., & Wilms, T. (2022). Effects of operational off-shore wind farms on fishes and fisheries. Review Report.

Thorstad, E. B., Todd, C. D., Uglem, I., Bjørn, P. A., Gargan, P. G., Vollset, K. W., Halttunen, E., Kålås, S., Berg, M., & Finstad, B. (2016). Marine life of the sea trout. Marine Biology, 163(3).

Thorstad, E. B., Whoriskey, F., Uglem, I., Moore, A., Rikardsen, A. H., & Finstad, B. (2012). A critical life stage of the Atlantic salmon Salmo salar: Behaviour and survival during the smolt and initial post-smolt migration. Journal of Fish Biology, 81(2).

Venables, W. N., & Ripley, B. D. (2002). Modern applied statistics with S. In Statistics and computing. Springer.

Verhelst, P., Pauwels, I., Pohl, L., Reubens, J., Schilt, B., & Hermans, A. (2025). Electromagnetic fields and diadromous fish spawning migration: An urgent call for knowledge. Marine Environmental Research, 204, 106857.

Williamson, B. J., Goddijn-Murphy, L., McIlvenny, J., & Youngson, A. (2024). Potential Exposure of Aquatic Organisms to Dynamic Visual Cues Originating from Aerial Wind Turbine Blades. Fishes, 9(12).

Wyman, M. T., Peter Klimley, A., Battleson, R. D., Agosta, T. V., Chapman, E. D., Haverkamp, P. J., Pagel, M. D., & Kavet, R. (2018). Behavioral responses by migrating juvenile salmonids to a subsea high-voltage DC power cable. Marine Biology, 165(8).