A. Gimpel et al. 
che captain and crew of the FFS Solea and the Team of the HC428 und 
SB765 surveys for their great support and flexibility. The stable isotope 
analysis was done at the Stable Isotope Analysis Laboratory of the Thünen 
Institute and supported by K. Hünerlage, thank you for great input. R. 
?rimicerio and M. H. Taylor is thanked for statistical guidance. We further 
:hank S. Duncan for advice. A.G. was funded under the European Maritime 
and Fisheries Fund of the European Union (partly EU Reg. 508/2014 and 
31S 989/AUF-FUEV/2-1/2018 “Co-Nutzung von Offshore-Windparks als 
ökosystembasiertes Raumplanungsmodell“). 
References 
Adlerstein, S.A., Ehrich, S., 2002, Effect of deviations from target speed and of time of day on 
zatch rates of some abundant species under North Sea International Bottom Trawl Survey 
protocol conditions. ICES J. Mar. Sci. 59, 594-603, 
Amoroso, R.O., Pitcher, C.R., Rijnsdorp, A.D., McConnaughey, R.A., Parma, A.M., Suuronen, 
P,, et al., 2018, Bottom trawl fishing footprints on the world's continental sheives, Proc. 
Natl. Acad. Sci. U. S. A. 115, E10275-E10282. 
Armstrong, M.J., Gerritsen, H.D., Allen, M., McCurdy, W.J., Peel, J.A.D., 2004. Variability in 
naturity and growth in a heavily exploited stock: cod (Gadus morhua L.) in the Irish Sea, 
ICES J. Mar. Sci. 61, 98-112. 
Aurich, H.J., 1941. Die Verbreitung der pelagischen Fischbrut in der südlichen Nordsee 
während der Frühjahrsfahrten 1926 der deutschen Forschungsschiffe, Poseidon” und, 
Makrele". Helgoländer wissenschaftliche Meeresuntersuchungen, N.F, 2, pp. 183-225 
1367 pp. 
Barbut, L., Vastenhoud, B., Vigin, L., Degraer, S., Volckaert, F.A.M., Lacroix, G., 2020. The pro- 
2ortion of flatfish recruitment in the North Sea potentially affected by offshore 
windfarms. ICES J. Mar. Sci. 77, 1227-1237. 
3ergström, L., Sundqvist, F., Bergström, U., 2013. Effects of an offshore wind farm on tempo- 
ral and spatial patterns in the demersal fish community. Mar. Ecol, Prog. Ser. 485, 
199-210. 
Callies, U., PIüß, A., Kappenberg, J., Kapitza, H., 2011. Particle tracking in the vicinity of 
Helgoland, North Sea: a model comparison. Ocean Dyn. 61, 2121-2139, 
Callies, U., Kreus, M., Petersen, W., Voynova, Y.G., 2021. On using Lagrangian drift simula- 
äons to aid interpretation of in situ monitoring data. Front. Mar. Sci, 8, 666653. 
https://doiorg/10.3389/fmars, 2021.666653, 
Campana, S.E., 2004. Photographic Atlas of Fish Otoliths of the Northwest Atlantic Ocean. 
NRC Research Press, Ottawa, 
Casini, M., Eero, M., Carlshamre, S., Lövgren, J., 2016. Using alternative biological informa- 
ion in stock assessment: condition-corrected natural mortality of Eastern Baltic cod. 
ICES J. Mar. Sci. 73, 2625-2631. 
Christiansen, N., Daewel, U., Djath, B., Schrum, C., 2022, Emergence of large-scale hydrody- 
namic structures due to atmospheric offshore wind farm wakes. Front. Mar. Sci. 9. 
Couce, E., Schratzberger, M., Engelhard, G.H., 2020. Reconstructing three decades of total in- 
ternational trawling effort in the North Sea. Earth Syst.Sci,.Data 12, 373-386, 
Jean, M.J., Hoffman, W.S., Zemeckis, D.R., Armstrong, M.P., 2014. Fine-scale diel and 
zender-based patterns in behaviour of Atlantic cod (Gadus morhua) on a spawning ground 
ın the Western Gulf of Maine. ICES J. Mar. Sci. 71 (6), 1474-1489. 
Degraer, S., Carey, D.A., Coolen, J.W.P., Hutchison, Z.L., Kerckhof, F., Rumes, B., et al., 2020. 
DIffshore wind farm artificial reefs affect ecosystem structure and functioning: a synthesis. 
Oceanography 33, 48-57. 
Dick, S., Kleine, E., 2007. The BSH new operational circulation model using general vertical 
co-ordinates. Environ.Res.Eng.Manag. 3, 18-24. 
Zhrich, S., Adlerstein, S., Brockmann, U., et al., 2007. 20 years of the German small-scale bot- 
tom trawi survey (GSBTS): a review. Senckenberg. Marit. 37, 13-82. 
ZU, 2018. European Union. Directive 2018/2001, revised Directive of the European Parlia- 
ment and of the Council of 11 December 2018 on the promotion of the use of energy 
from renewable sources, https://eur-lex.europa.eu/eli/dir/2018/2001/0j. 
70x, C.J., Taylor, M., Dickey-Collas, M., Fossum, P., Kraus, G., Rohlf, N., et al., 2008. Mapping 
he spawning grounds of North Sea cod (Gadus morhua) by direct and indirect means, 
Proc. R. Soc. B Biol. Sci. 275, 1543-1548. 
Gimpel, A., Stelzenmüller, V., Cormier, R., Floeter, J., Temming, A., 2013. A spatially explicit 
:isk approach to support marine spatial planning in the German EEZ, Mar. Environ. Res. 
86, 56-69. 
Gourvenec, S., Sturt, F., Reid, E., Trigos, F., 2022, Global assessment of historical, current and 
‘orecast ocean energy infrastructure: implications for marine space planning, sustainable 
design and end-of-engineered-life management. Renew. Sust. Energ, Rev. 154. 
Greenacre, M., Primicerio, R., 2013. Multivariate Analysis of Ecological Data. BBVA Founda- 
tion, Bilbao. 
Guiet, J., Galbraith, E., Kroodsma, D., Worm, B., 2019. Seasonal variability in global industrial 
fishing effort. PLoS ONE 14. 
Gusatu, L.F., Menegon, S., Depellegrin, D., Zuidema, C., Faaij, A., Yamı, C., 2021. Spatial and 
‚emporal analysis of cumulative environmental effects of offshore wind farms in the 
North Sea basin. Sci. Rep. 11, 10125, 
Aaggarty, D.R., King, J.R., 2006. CPUE as an index of relative abundance for nearshore reef 
fishes. Fish. Res. 81. 
Jammar, L., Wikström, A., Molander, S., 2014. Assessing ecological risks of offshore wind 
power on Kattegat cod, Renew. Energy 66, 414-424. 
Hayward, P.J., Ryland, J.S., 1990a. The marine fauna of the British Isles and North-West 
Europe. Introduction And Protozoans to Arthropods. Vol. 1. Clarendon Press. 
Hayward, P.J., Ryland, J.S., 1990b. The marine fauna of the British Isies and North-West 
Europe. Molluscs to ChordatesVol.2. Clarendon Press. 
Science of the Total Environment 878 (2023) 162902 
Aeath, M.R., Kunzlik, P.A., Gallego, A., Holmes, S.J., Wright, P.J., 2008. A model of meta- 
Dopulation dynamics for North Sea and West of Scotland cod—the dynamic consequences 
of natal fidelity. Fish. Res. 93, 92-116. 
Aünerlage, K., Buchholz, F., 2013. Krill of the northern Benguela Current and the Angola-Ben- 
zuela frontal zone compared: physiological performance and short-term starvation in 
£uphausia hanseni. J. Plankton Res. 35, 337-351. 
CES, 2014. Report of the Workshop on Sexual Maturity Staging of Cod, Whiting, Had-dock, 
Saithe And Hake (WKMSGAD), 4-8 November 2013, San Sebastian, Spain. ICES CM 
2013\ACOM:57 51 pp. 
CES, 2022. Working Group on the Assessment of Demersal Stocks in the North Sea and Skag- 
arrak (WGNSSK). ICES Scientific Reports. 4, p. 43 1367 pp. 
Gjesbu, O., Klungseyr, J., Kryvi, H., Witthames, P.W., Walker, M.G., 1991. Fecundity, atresia 
and egg size of captive Atlantic Cod (Gadus morhua) in relation to proximate body com- 
position, Can. J. Fish, Aquat. Sci. 48, 2333-2343. 
{raufvelin, P., Bergstrom, L., Sundqvist, F., Ulmestrand, M., Wennhage, H., Wikstrom, A., et 
al., 2023. Rapid re-establishment of top-down control at a no-take artificial reef. Ambio 
52, 556-570. 
<urlansky, M., 1997. Cod: A Biography of the Fish That Changed the World. Walker and Co., 
New York, p. 294. 
„anghamer, O., 2012. Artificial reef effect in relation to offshore renewable energy conver- 
sion: state of the art. ScientificWorldJournal 2012, 386713. 
‚eonhard, S.B., Stenberg, C., Stettrup, J., Deurs, M.V., Christensen, A., Pedersen, J., 2013. Fish 
benefits from offshore wind farm development. Danish Offshore Wind - Key Environmen- 
cal Issues - A Follow-up. Danish Energy Agency, Copenhagen, pp. 31-45. 
„ewis, L.A., Richardson, D.E., Zakharov, E.V., Hanner, R., 2016. Integrating DNA barcoding of 
fish eggs into ichthyoplankton monitoring programs. Fish. Bull. 114, 153-165. 
‚indeboom, H., Degraer, S., Dannheim, J., Gill, A.B., Wilhelmsson, D., 2015. Offshore wind 
Dark monitoring programmes, lessons learned and recommendations for the future. 
Hydrobiologia 756, 169-180. 
Vünez-Riboni, I., Taylor, M.H., Kempf, A., Püts, M., Mathis, M., Ojaveer, H., 2019. Spatially 
resolved past and projected changes of the suitable thermal habitat of North Sea cod 
(Gadus morhua) under climate change. ICES J. Mar. Sci. 76, 2389-2403. 
Ovegärd, M., Berndt, K., Lunneryd, S.G., 2012. Condition indices of Atlantic cod (Gadus 
morhua) biased by capturing method. ICES J. Mar. Sci. 69, 1781-1788. 
?»örtner, H.O., Karl, D.M., Boyd, P.W., Cheung, W.W.L., Liuch-Cota, S.E., Nojiri, Y., et al., 
2014. Ocean systems. In: Field, C.B., Barros, V.R., Dokken, D.J., Mach, K.J., 
Mastrandrea, M.D., Bilir, T.E., Chatterjee, M., Ebi, K.L., Estrada, Y.O., Genova, R.C., 
Sirma, B., Kissel, E.S., Levy, A.N., MacCracken, S., Mastrandrea, P.R., White, L.L. (Eds.), 
Zlimate Change 2014: Impacts, Adaptation, And Vulnerability, Part A: Global And Sec- 
:oral Aspects, Contribution of Working Group II to the Fifth Assessment Report of the In- 
;ergovernmental Panel on Climate Change. Cambridge University Press, Cambridge, 
United Kingdom and New York, NY, USA, pp. 411484. 
Reubens, J., Degraer, S., Vincx, M., 2012. The secret life of Atlantic cod (Gadus morhua) at a 
wind farm in the Beigian part of the North Sea: where ecology meets economy. In: Mees, 
J., et al. (Eds.), Book of Abstracts - VLIZ Young Marine Scientists' Day. Brugge, Belgium, 
24 February 2012. VLIZ Special Publication. 55, p. 69. 
Reubens, J., Pasotti, F., Degraer, S., Vincx, M., 2013a. Residency, site fidelity and habitat use 
of Atlantic cod (Gadus morhua) at an offshore wind farm using acoustic telemetry. Mar. 
Environ. Res, 90, 128-135. 
Reubens, J.T., Braeckman, U., Vanaverbeke, J., Van Colen, C., Degraer, S., Vincx, M., 2013b. 
Aggregation at windmiill artificial reefs: CPUE of Atlantic cod (Gadus morhua) and 
pouting (Trisopterus hıscus) at different habitats in the Belgian part of the North Sea. 
Fish. Res. 139, 28-34. 
Reubens, J.T., Vandendriessche, S., Zenner, A.N., Degraer, S., Vincx, M., 2013c. Offshore wind 
“arms as productive sites or ecological traps for gadoid fishes?-impact on growth, condi- 
tion index and diet composition. Mar. Environ. Res. 90, 66-74. 
Zenbens, J.T., De Riüjcke, M., Degraer, S., Vincx, M., 2014a. Diel variation in feeding and move- 
ment patterns of juvenile Atlantic cod at offshore wind farms, J, Sea Res. 85, 214-221. 
zeubens, J.T., Degraer, S., Vincx, M., 2014b. The ecology of benthopelagic fishes at offshore 
wind farms: a synthesis of 4 years of research. Hydrobiologia 727, 121-136. 
Rossel, S., Barco, A., Kloppmann, M.H.F., Arbizu, P.M., Huwer, B., Knebelsberger, T., 2021. 
Rapid species level identification of fish eggs by proteome fingerprinting using MALDI- 
TOF MS. J. Proteome 231. 
Zussell, F.S., 1976. The Eggs And Planktonic Stages of British Marine Fishes. Academic Press 
524 pp. 
Russell, D.J.F., Brasseur, S., Thompson, D., Hastie, G.D., Janik, V.M., Aarts, G., et al., 2014. 
Marine mammals trace anthropogenic structures at sea. Curr. Biol, 24, R638-R639. 
Stelzenmüller, V., Fock, H.O., Gimpel, A., Seidel, H., Diekmann, R., Probst, W.N., et al., 2015, 
Quantitative environmental risk assessments in the context of marine spatial manage- 
ment: current approaches and some perspectives. ICES J. Mar. Sci. 72 (3), 1022-1042. 
nttps://doi.org/10.1093/icesjms/fsu206. 
Stelzenmüller, V., Letschert, J., Gimpel, A., Kraan, C., Probst, W.N., Degraer, S., et al., 2022. 
From plate to plug: the impact of offshore renewables on European fisheries and the 
role of marine spatial planning. Renew, Sust. Energ. Rev. 158. 
'hompson, B.M., Riley, J.D., 1981. Egg and jarvail development studies in the North Sea Cod 
(Gadus morhua). Rapp. P.-v. Reun. Cons. int. Explor, Mer 178, 553-559. 
zan Berkel, J., Burchard, H., Christensen, A., Mortensen, L.O., Svenstrup Petersen, O., 
"homsen, F., 2020. The effects of offshore wind farms on hydrodynamics and implica- 
cions for fishes. Oceanography 33 (4), 108-117. 
Vitale, F., Börjesson, P., Svedäng, H., Casini, M., 2008. The spatial distribution of cod (Gadus 
morhua L.) spawning grounds in the Kattegat, eastern North Sea, Fish. Res. 90, 36-44, 
Werner, K.M., Taylor, M.H., Diekmann, R., Lloret, J., Möllmann, C., Primicerio, R., et al., 
2019, Evidence for limited adaptive responsiveness to large-scale spatial variation of hab- 
itat quality. Mar. Ecol. Prog. Ser, 629, 179-191. 
Zimmermann, F., Werner, K.M., 2019. Improved management is the main driver behind re- 
coverv of Northeast Atlantic fish stocks. Front. Ecol. Environ. 17, 93-99