24While considerable effort has been devoted to understanding the factors regulating the 25 development of phytoplankton blooms, the mechanisms leading to bloom decline and 26 termination have received less attention. Grazing and sedimentation have been invoked as 27 the main routes for the loss of phytoplankton biomass, and more recently, viral lysis, 28 parasitism and programmed cell death (PCD) have been recognized as additional removal 29 factors. Despite the importance of bloom declines to phytoplankton dynamics, the 30 incidence and significance of various loss factors in regulating phytoplankton populations 31have not been widely characterized in natural blooms. To understand mechanisms 32 controlling bloom decline, we studied two independent, inshore blooms of Alexandrium 33 fundyense, paying special attention to cell mortality as a loss pathway. We observed 34 increases in the number of dead cells with PCD features after the peak of both blooms, 35 demonstrating a role for cell mortality in their terminations. In both blooms, sexual cyst 36 formation appears to have been the dominant process leading to bloom termination, as 37 both blooms were dominated by small-sized gamete cells near their peaks. Cell death and 38 parasitism became more significant as sources of cell loss several days after the onset of 39 bloom decline. Our findings show two distinct phases of bloom decline, characterized by 40 sexual fusion as the initial dominant cell removal processes followed by elimination of 41 remaining cells by cell death and parasitism. factors (Cloern 1996). Blooms occur when growth processes are dominant over losses, 47 then decline when loss processes intensify relative to growth. The mechanisms 48 underlying loss processes are diverse and poorly constrained. Traditionally, grazing by 49 heterotrophic zooplankton and sinking into the deep ocean have been considered the main 50 loss factors for phytoplankton (Calbet and Landry 2004). However, natural populations 51 and laboratory cultures often collapse abruptly due to massive cell lysis events, indicating 52 that different processes must be contributing to phytoplankton mortality (Berges and 53 Falkowski 1998;Vardi et al. 1999; Berges and Choi 2014). Virus-induced cell lysis is 54 increasingly recognized as an important alternative loss factor, with a large number of 55 viruses found in aquatic ecosystems (Brussaard 2004). There is evidence that chronic 56 parasitic infections can be an important cause of phytoplankton population crashes as 57 well (Chambouvet et al. 2008). 58Another loss mechanism that has received increased attention in recent years is 59 autocatalyzed cell mortality, such as programmed cell death (PCD). PCD is different 60 from other pathways of cell death because it is an active process that is initiated by the 61 cell itself and tightly controlled by gene expression and protein synthesis inside of the 62 cells (Bidle and Falkowski 2004). Accumulating evidence suggests that PCD occurs in 63 phytoplankton under diverse envi...