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. Coupled physical–biological models usually resolve only parts of the trophicfood chain; hence, they run the risk of neglecting relevant ecosystemprocesses. Additionally, this imposes a closure term problem at therespective “ends” of the trophic levels considered. In this study, we aim tounderstand how the implementation of higher trophic levels in anutrient–phytoplankton–zooplankton–detritus (NPZD) modelaffects the simulated response of the ecosystem using a consistentNPZD–fish modelling approach (ECOSMO E2E) in the combined North Sea–Baltic Sea system.Utilising this approach, we addressed the above-mentionedclosure term problem in lower trophic ecosystem modelling at a very lowcomputational cost; thus, we provide an efficient method that requires verylittle data to obtain spatially and temporally dynamic zooplankton mortality. On the basis of the ECOSMO II coupled ecosystem model we implemented onefunctional group that represented fish and one group that represented macrobenthosin the 3-D model formulation. Both groups were linked to the lower trophiclevels and to each other via predator–prey relationships, which allowed for theinvestigation of both bottom-up processes and top-down mechanisms in thetrophic chain of the North Sea–Baltic Sea ecosystem. Model results for a10-year-long simulation period (1980–1989) were analysed and discussed withrespect to the observed patterns. To understand the impact of the newlyimplemented functional groups for the simulated ecosystem response, wecompared the performance of the ECOSMO E2E to that of a respective truncatedNPZD model (ECOSMO II) applied to the same time period. Additionally, weperformed scenario tests to analyse the new role of the zooplankton mortalityclosure term in the truncated NPZD and the fish mortality term in theend-to-end model, which summarises the pressure imposed on the system byfisheries and mortality imposed by apex predators. We found that the model-simulated macrobenthos and fish spatial and seasonalpatterns agree well with current system understanding. Considering a dynamicfish component in the ecosystem model resulted in slightly improved modelperformance with respect to the representation of spatial and temporal variationsin nutrients, changes in modelled plankton seasonality, and nutrient profiles.Model sensitivity scenarios showed that changes in the zooplankton mortalityparameter are transferred up and down the trophic chain with littleattenuation of the signal, whereas major changes in fish mortality and fishbiomass cascade down the food chain.

Towards end-to-end (E2E) modelling in a consistent NPZD-F modelling framework (ECOSMO E2E_v1.0): application to the North Sea and Baltic Sea