Why do red tides occur frequently in some oligotrophic waters? Analysis of red tide evolution history in Mirs Bay, China and its implications.

The process and management of red tide in oligotrophic waters are poorly understood as most studies on red tide were focused on eutrophic areas. In this study, 404 red tide events together with the historical water quality dynamics during 1991-2020 were investigated in an anthropogenically influenced bay in China - Mirs Bay, whose most region is oligotrophic except small inshore areas. Red tides of oligotrophic offshore accounted for 20 % of all. With the effective governmental management on inshore areas, concentration of PO4 and DIN has been decreased to a low level (PO4 <0.01 mg/L while DIN <0.1 mg/L) in the bay since about 2000. However, the reduction of nutrients was still accompanied by the frequent outbreaks of red tides, as well as a shift of dominant algae from diatoms to dinoflagellates, which might be due to the unbalanced nutrient reduction, such as N:P ratio fluctuation and organic nutrient increase. This shift might trigger more red tide events and even some super ones (long-duration or large-scale) in oligotrophic areas. Detailed analysis on red tide events combined with model simulation proved that the outbreak of red tide in Mirs Bay was caused by the joint contribution of nutrients and hydrodynamics. Nutrients of inshore area supported the red tides there, and with the help of physical conditions, red tides inshore could be transferred to offshore areas and then were likely to bloom again or be preyed to support blooms of other organisms. This study acknowledged that the reduction of both N and P either inorganic or organic nutrients was essential to control red tides, even in oligotrophic waters, but a balanced strategy considering the dual reduction of both nitrogen and phosphorus was of pivotal role to restore the health of coastal water systems disturbed by human.