Ecological features of harmful algal blooms in coastal upwelling ecosystems

Upwelling regions are the most complex habitats in which dinoflagellates product red tides, but the flora is not unique. Many species also bloom in nutrient-enriched, non-upwelling systems, share the collective dinoflagellate trait of low-nutrient affinity, and can achieve relatively fast growth rates. Blooms occur over the range of nutrient - mixing - advection combinations found in upwelling habitats, rather than bring restricted to the high-nutrient high-irradiance low-turbulence conditions posited by Margalef's classical Mandala and its Bowman rr al. and Pingree versions. The bloom species are primarily ruderal strategists (R-species), which typify "mixing - drift" life-forms adapted to the velocities associated with frontal zones, entrainment within coastal currents, and vertical mixing during upwelling relaxations. Collectively, dinoflagellates appear capable of surviving fairly high turbulence spectra formed at representative Kolmogorov length scale - wind speed conditions. This biophysical protection might be the result of cell size-facilitated entrainment within the micro-eddies formed during turbulent energy dissipation. The swimming speeds of 71 clones of dinoflagellates are compared and related to reported rates of vertical motion in coastal upwelling systems. There are slow and fast swimmers: many exhibit motility rates that can exceed representative in situ vertical and horizontal water mass movements. At least four dinoflagellates from upwelling systems form chains leading to increased swimming speeds, and may be an adaptation for growth in coastal upwelling habitats. Red tides are frequent and fundamental features of upwelling systems, particularly during intermittent upwelling relaxations, rather than dichotomous (sometimes catastrophic) interruptions of the diatom blooms characteristically induced by upwelling. Successional sequences and the "red tide" zone may differ between upwelling and non-upwelling systems. In the latter, red tides diverge from the main sequence and are appropriately positioned in the Mandala's ecological space of high nutrients and low turbulence. An amended Mandala based on Pingree's S-kh model and the Smayda and Reynolds life-form model is presented to accommodate the range of red tide development and their successional routing found in coastal upwelling systems. Ecophysiological data support the pitcher and Boyd seeding mechanism model, which can lead to red tides in upwelling systems. Nutrients, phyto-stimulation and grazing pressure as triggering factors in upwelling-system red tides are considered. Some red tides may be stimulated by nutrients and growth promoting factors excreted by migrating shoals and "boils" of clupeoid stocks, with selective zoo-plankton grazing contributory. Substantial collapses in grazing pressure may be essential in anoxic red tide events. The mass mortalities that accompany anoxia, common to the Benguela and Peru upwelling systems, may be a trophic control mechanism to maintain biogeochemical balance and regional homeostasis, which are vital to upwelling ecosystem dynamics. Some traditional concepts of phytoplankton ecology may not completely apply to dinoflagellate bloom events in coastal upwelling systems.