Dinoflagellate blooms in upwelling systems: Seeding, variability, and contrasts with diatom bloom behaviour

The influence of diatom bloom behaviour, dinoflagellate life cycles, propagule type and upwelling bloom cycles on the seeding of dinoflagellate blooms in eastern boundary current upwelling systems is evaluated. Winter-spring diatom bloom behaviour is contrasted with upwelling bloom behaviour because their phenology impacts dinoflagellate blooms. The winter-spring diatom bloom is usually sustained, whereas the classical upwelling diatom bloom occurs as a series of separate, recurrent mini-blooms intercalated by upwelling-relaxation periods, during which dinoflagellates often bloom. Four sequential wind-regulated phases characterize upwelling cycles, with each phase having different effects on diatom and dinoflagellate bloom behaviour: bloom spin up, bloom maximum, bloom spin down, and upwelling relaxation. The spin up - bloom maximum is the period of heightened diatom growth; the spin down - upwelling-relaxation phases are the periods when dinoflagellates often bloom. The duration, intensity and ratio of the upwelling and relaxation periods making up upwelling cycles determine the potential for dinoflagellate blooms to develop within a given upwelling cycle and prior to the subsequent spin up of upwelling that favours diatom blooms. Upwelling diatoms and meroplanktonic dinoflagellates have three types of propagules available to seed blooms: vegetative cells, resting cells and resting cysts. However, most upwelling dinoflagellates are holoplanktonic, which indicates that the capacity to form resting cysts is not an absolute requirement for growth and survival in upwelling systems. The long-term (decadal) gaps in bloom behaviour of Gymnodinium catenatum and Lingulodinium polyedrum, and the unpredictable bloom behaviour of dinoflagellates generally, are examined from the perspective of seeding strategies. Mismatches between observed and expected in situ bloom behaviour and resting cyst dynamics are common among upwelling dinoflagellates. This disassociation suggests unrecognized upwelling system factors that fall within the physical-chemical-biological domain are more important than life cycle in selecting dinoflagellates species having the survival-seeding strategies and ecophysiological adaptations required for growth in physically robust upwelling systems. It is conjectured that diatom life cycles, as a group, are geared towards exploiting seeding opportunities, whereas dinoflagellates have evolved life-cycle behaviour more attuned to survival. The role of ecological dormancy and ecological release from bloom inhibition underlying dinoflagellate bloom irregularity is considered. The expectation that the dinoflagellate species selected to bloom from among the common upwelling flora would be the same in all eastern boundary upwelling systems is not realized. (C) 2010 Elsevier Ltd. All rights reserved.