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A coupled water-quality–Ecopath with Ecosim model incorporating sand-lance aestivation dynamics in the Harima-Nada Sea

Year: 2026
Type: article
Abstract: Sand lance ( Ammodytes japonicus ) is an important forage fish in the Harima-Nada Sea, Japan, but its stock has declined markedly in recent decades under concurrent changes in nutrient availability and water temperature. To clarify the mechanisms underlying these changes, we developed a region-specific coupled water-quality–Ecopath with Ecosim (EwE) model that explicitly incorporates sand-lance aestivation dynamics. The model integrates temperature-dependent production and consumption, nutrient-dependent primary production, an empirical relationship between nutrient levels and dissolved oxygen, DO-dependent non-predation mortality, and temperature-dependent onset, termination, and mortality during sand-lance aestivation. Model performance was evaluated using PREBAL diagnostics and comparisons with historical biomass, catch, and phytoplankton data. We then conducted single-driver sensitivity analyses for nutrient concentration, water temperature, prey availability, and predator biomass to isolate the mechanistic pathways through which each driver affects sand-lance biomass. The model reproduced major long-term patterns in the Harima-Nada food web and indicated that sand-lance biomass was strongly controlled by bottom-up processes mediated through nutrient availability and Calanus sinicus . Moderate nutrient enrichment enhanced sand-lance biomass by supporting lower-trophic production, whereas excessive enrichment increased hypoxia-related mortality and weakened trophic transfer. Warming reduced sand-lance biomass by prolonging the aestivation period, shortening the productive season, increasing high-temperature mortality during aestivation, and reducing the availability of thermally sensitive prey. Under the present scenario design, prey-availability scenarios produced larger biomass responses than predator-biomass scenarios, indicating a particularly strong bottom-up pathway in the model. These results demonstrate that incorporating species-specific aestivation dynamics into EwE provides a mechanistic basis for evaluating sand-lance responses to environmental change and for developing future multi-driver applications in coastal ecosystem management.
Language: English
Open Access status: hybrid
Estimated APC paid: $2,810
OpenAlex ID: W7218694630
FWCI:
Cites: 27
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