Complex trophic interactions of calanoid copepods in the Benguela upwelling system

Life-cycle adaptations, dietary preferences and trophic levels of calanoid copepods from the northern Benguela Current off Namibia were determined via lipid classes, marker fatty acids and stable isotope analyses, respectively. Trophic levels of copepod species were compared to other zooplankton and top consumers. Lipid class analyses revealed that three of the dominant calanoid copepod species stored wax esters, four accumulated triacylglycerols and another three species were characterised by high phospholipid levels. The two biomarker approaches (via fatty acids and stable isotopes) revealed a complex pattern of trophic positions for the various copepod species, but also highlighted the dietary importance of diatoms and dinoflagellates. Calanoides carinatus and Nannocalanus minor occupied the lowest trophic level (predominantly herbivorous) corresponding to high amounts of fatty acid markers for diatoms (e.g. 16:1(n − 7)) and dinoflagellates (e.g. 18:4(n − 3)). These two copepod species represent the classical link between primary production and higher trophic levels. All other copepods belonged to secondary or even tertiary (some deep-sea copepods) consumers. The calanoid copepod species cover the entire range of δ15N ratios, as compared to δ15N ratios of all non-calanoid taxa investigated, from salps to adult fish. These data emphasise that the trophic roles of calanoid copepods are far more complex than just interlinking primary producers with pelagic fish, which should also be considered in the process of developing realistic food-web models of coastal upwelling systems.