The role of nutrients in regulation and promotion of harmful algal blooms in upwelling systems

The Core Research Project on HABs in upwelling systems, as a component project of the international scientific programme on the Global Ecology and Oceanography of Harmful Algal Blooms (GEOHAB), promotes a comparative approach within and across systems to understand and predict the ecology, frequency and occurrence of HABs in eastern boundary current upwelling systems. Unlike other systems, upwelling circulation tends to override nutrient limitation caused by stratification, but is less affected by anthropogenic impacts due to the magnitude of the upwelling nutrient signal. At the same time, upwelling systems are unique in that they undergo seasonal succession as well as short-term spatial and temporal oscillations driven by the time-scale of upwelling wind events. An understanding of nutrient dynamics in upwelling systems is thus critical to any attempt to understand or predict HAB events in these environments. We review the state of knowledge regarding nutrient ecophysiology of a subset of HAB organisms identified in upwelling systems. The upwelling HABs exhibit a number of adaptations previously identified in HAB organisms, such as mixotrophy, osmotrophy and vertical migration. We suggest that, unlike most other HABs, these organisms do not necessarily follow a low nutrient-affinity strategy, and do not fit well with classic allometric scaling relationships. Despite these anomalies, progress has been made in predicting HAB events in upwelling systems, by linking HAB events to the unique environmental conditions associated with these systems. We conclude that this subset of HAB organisms is still poorly described in terms of nutrient ecophysiology, and will benefit from a comparative approach across systems, particularly because the subset of upwelling HABs does not necessarily fit the generic patterns identified for HABs generally.