The physical oceanography of upwelling systems and the development of harmful algal blooms.

The upwelling systems of the eastern boundaries of the world's oceans are susceptible to harmful algal blooms (HABs) because they are highly productive, nutrient-rich environments, prone to high-biomass blooms. This review identifies those aspects of the physical environment important in the development of HABs in upwelling systems through description and comparison of bloom events in the Benguela, California and Iberia systems. HAB development is dictated by the influence of wind stress on the surface boundary layer through a combination of its influence on surface mixed-layer characteristics and shelf circulation patterns. The timing of HABs is controlled by windstress fluctuations and buoyancy inputs at the seasonal, event and interannual scales. Within this temporal framework, various mesoscale features that interrupt typical upwelling circulation patterns, determine the spatial distribution of HABs. The inner shelf in particular provides a mosaic of shifting habitats, some of which favour HABs. Changes in coastline configuration and orientation, and bottom topography are important in determining the distribution of HABs through their influence on water stratification and retention. A spectrum of coastline configurations, including headlands, capes, peninsulas, Rías, bays and estuaries, representing systems of increasing isolation from the open coast and consequent increasing retention times, are assessed in terms of their vulnerability to HABs.