Description: Euphausiids constitute major biomass component in shelf ecosystems and play a fundamental role in the rapid vertical transport of carbon from the ocean surface to the deeper layers during their daily vertical migration (DVM). DVM depth and migration patterns depend on oceanographic conditions with respect to temperature, light and oxygen availability at depth, factors that are highly dependent on season in most marine regions. Changes in the abiotic conditions also shape Euphausiid metabolism including aerobic and anaerobic energy production. Here we introduce a global krill respiration model which includes the effect of latitude (LAT), the day of the year of interest (DoY), and the number of daylight hours on the day of interest (DLh), in addition to the basal variables that determine ectothermal oxygen consumption (temperature, body mass and depth) in the ANN model (Artificial Neural Networks). The newly implemented parameters link space and time in terms of season and photoperiod to krill respiration. The ANN model showed a better fit (r**2=0.780) when DLh and LAT were included, indicating a decrease in respiration with increasing LAT and decreasing DLh. We therefore propose DLh as a potential variable to consider when building physiological models for both hemispheres. We also tested for seasonality the standard respiration rate of the most common species that were investigated until now in a large range of DLh and DoY with Multiple Linear Regression (MLR) or General Additive model (GAM). GAM successfully integrated DLh (r**2= 0.563) and DoY (r**2= 0.572) effects on respiration rates of the Antarctic krill, Euphausia superba, yielding the minimum metabolic activity in mid-June and the maximum at the end of December. Neither the MLR nor the GAM approach worked for the North Pacific krill Euphausia pacifica, and MLR for the North Atlantic krill Meganyctiphanes norvegica remained inconclusive because of insufficient seasonal data coverage. We strongly encourage comparative respiration measurements of worldwide Euphausiid key species at different seasons to improve accuracy in ecosystem modelling.

Description: Euphausiids constitute a major biomass component in shelf ecosystems and play a fundamental role in the rapid vertical transport of carbon from the ocean surface to the deeper layers during their daily vertical migration (DVM). DVM depth and migration patterns depend on oceanographic conditions with respect to temperature, light and oxygen availability at depth, factors that are highly dependent on season in most marine regions. Here we introduce a global krill respiration ANN (artificial neural network) model including the effect of latitude (LAT), the day of the year (DoY), and the number of daylight hours (DLh), in addition to the basal variables that determine ectothermal oxygen consumption (temperature, body mass and depth). The newly implemented parameters link space and time in terms of season and photoperiod to krill respiration. The ANN model showed a better fit (r2 = 0.780) when DLh and LAT were included, indicating a decrease in respiration with increasing LAT and decreasing DLh. We therefore propose DLh as a potential variable to consider when building physiological models for both hemispheres. For single Euphausiid species investigated in a large range of DLh and DoY, we also tested the standard respiration rate for seasonality with Multiple Linear Regression (MLR) and General Additive model (GAM). GAM successfully integrated DLh (r2 = 0.563) and DoY (r2 = 0.572) effects on respiration rates of the Antarctic krill, Euphausia superba, yielding the minimum metabolic activity in mid-June and the maximum at the end of December. We could not detect DLh or DoY effects in the North Pacific krill Euphausia pacifica, and our findings for the North Atlantic krill Meganyctiphanes norvegica remained inconclusive because of insufficient seasonal data coverage. We strongly encourage comparative respiration measurements of worldwide Euphausiid key species at different seasons to improve accuracy in ecosystem modeling.

Description: Euphausiids constitute a major biomass component in shelf ecosystems and play a fundamental role in the rapid vertical transport of carbon from the ocean surface to the deeper layers during their daily vertical migration (DVM). DVM depth and migration patterns depend on oceanographic conditions with respect to temperature, light and oxygen availability at depth, factors that are highly dependent on season in most marine regions. Here we introduce a global krill respiration ANN (Artificial Neural Network) model including the effect of latitude (LAT), the day of the year (DoY), and the number of daylight hours (DLh), in addition to the basal variables that determine ectothermal oxygen consumption (temperature, body mass and depth). The newly implemented parameters link space and time in terms of season and photoperiod to krill respiration. The ANN model showed a better fit (r2=0.780) when DLh and LAT were included, indicating a decrease in respiration with increasing LAT and decreasing DLh. We therefore propose DLh as a potential variable to consider when building physiological models for both hemispheres. For single Euphausiid species investigated in a large range of DLh and DoY, we also tested the standard respiration rate for seasonality with Multiple Linear Regression (MLR) and General Additive model (GAM). GAM successfully integrated DLh (r2= 0.563) and DoY (r2= 0.572) effects on respiration rates of the Antarctic krill, Euphausia superba, yielding the minimum metabolic activity in mid-June and the maximum at the end of December. We could not detect DLh or DoY effects in the North Pacific krill Euphausia pacifica, and our findings for the North Atlantic krill Meganyctiphanes norvegica remained inconclusive because of insufficient seasonal data coverage. We strongly encourage comparative respiration measurements of worldwide Euphausiid key species at different seasons to improve accuracy in ecosystem modelling.

Description: In the framework of the GENUS –“Geochemistry and Ecology of the Namibian Upwelling System” research program, trophic interactions and carbon pathways throughout the food web of the coastal upwelling system are being quantified. In contrast to earlier studies, special focus is been given to lower trophic levels in higher taxonomic resolution. Energy demands of various zooplankton taxa, including copepods, euphausiids, decapods and fish larvae, have been quantified with standardized methodology via optode respirometry. Dietary spectra and trophic levels were analyzed by trophic biomarker approaches based on fatty acid composition and stable isotopes (15N, 13C), respectively. All empirical data are assembled for an Ecopath with Ecosim (EwE) food-web model. The EwE model distinguishes between shelf and offshore communities. The conceptual food‐web model consists of three groups of primary producers, i.e. diatoms, dinoflagellates, and cyanobacteria, as well as many consumers such as Calanoides carinatus as the key herbivorous copepod in the Benguela upwelling system, other copepods, Euphausia hanseni, other zooplankton, and pelagic fishes including sardine, anchovy, and horse mackerel. Empirical data show that zooplankton and particularly copepods encompass a wider range of trophic levels from herbivory to secondary or even tertiary consumers (δ15N from 4 to 12‰), while anchovy had rather low δ15N of about 7‰. Respiration rates and metabolic activities of copepods could be parameterized for the model by an energy budget approach based on ambient temperature, body mass, and activity level. Calanoid copepods consumed 78mg C m‐2 d‐1 in shelf regions and 21mg C m‐2 d‐1 in oceanic regions. Locally, C. carinatus could remove up to 90% of the diatom biomass per day. The community consumption of pelagic decapods ranged from 7 mg C m‐2 d‐1 to 〉20mg C m‐2 d‐1 with highest values in the northernmost part of the study area. Overall, pelagic decapods apparently play a more prominent role in the northern Benguela ecosystem than previously assumed and may exert a substantial predation pressure on calanoid copepods. GENUS results emphasize that the trophic interactions within zooplankton and lower trophic levels are more complex than just linking primary producers with pelagic fish and should be taken into account in the process of developing realistic food‐web models of coastal upwelling systems. Keywords: foodweb, zooplankton, trophic interactions, energy flux

Description: The goal of the GENUS project (Geochemistry and Ecology of the Namibian Upwelling System) is to analyse the interrelationships between climate change, oceanic nutrients, greenhouse gases and the ecosystem structure in the coastal upwelling area off Namibia. The biological/ecological work package focused on the structure of the Northern Benguela Upwelling System (NBUS) and its energy flows under changing environmental conditions. Physiological constraints and adaptations were detected in several taxonomic groups investigated within the project, such as several copepod species, euphausiids and fish larvae. Temperature and oxygen distribution in the water column were identified as main drivers in modulating the distribution and ecology of many species. The extension and position of the Oxygen Minimum Zone (OMZ) seems to have a significant impact on the life cycles, vertical distribution and trophic condition of various species. In copepods we find a variety of adaptation mechanisms. While some species avoid the OMZ, others use it for resting and predator avoidance during daytime. Such vertical migrations contribute significantly to vertical carbon flux. The same holds true for decapods (4.4 mg C m‐2 d‐1). Respiration rates of 16 copepod species were determined to average 54.6 ±32.8 ml O2 d‐1 gDM‐1. Calanoides carinatus diapausing C5 stages reduced respiration at depth by 82% compared to surface activity. Further adaptations were found in euphausiid species: While Euphausia hanseni is capable to use the OMZ as a retreat by reducing its metabolic activity at lower temperatures and unfavourable trophic conditions, Nematoscelis megalops generally maintains a low level metabolism adapted to a constant life in the OMZ, and avoids crossing the thermocline. Special features of early life stages of Trachurus capensis, the fish species actually showing highest commercial landings, were analysed to elucidate their potential advantages in life performance, compared to other small pelagic species such as sardines or anchovies. The species showed short‐term hypoxia tolerance down to 30% oxygen saturation and even survived 10% saturation. Combined with the ability to switch from smaller to larger copepod prey and to surpass vulnerable early stages much faster than competitor species, this could explain the dominance of Trachurus capensis in the NBUS. In addition, competitors and predators of fish larvae such as jellyfish or chaetognaths showed little or no response to low oxygen concentrations gaining advantage over e.g. sardines and anchovies. Isotope analyses of various pelagic species and their food revealed a complex picture of the trophic levels of species including developmental stages and provide the basis for trophic flow models. The results will also serve to calibrate carbon dynamics and nutrient flux models that were developed in another GENUS work package. Data will be compared with nutrient distribution patterns and dynamics that may influence primary production and impact zooplankton distribution and higher trophic levels such as fish, seabirds or mammals. Results clearly show that continuing ocean warming coupled with expansion of the OMZ may alter horizontal and vertical distribution of species and the food web structure of the ecosystem. Keywords: Benguela Current, OMZ, Pelagic Ecosystem, Physiology