Calcifying marine phytoplankton - coccolithophores - are some of the most successful yet enigmatic organisms in the ocean, and are at risk from global change. In order to better understand how they will be affected we need to know ‘why’ coccolithophores calcify. Here we review coccolithophorid evolutionary history, cell biology, and insights from recent experiments to provide a critical assessment of the costs and benefits of calcification. We conclude that calcification has high energy demands, and that coccolithophores might have calcified initially to reduce grazing pressure, but that additional benefits such as protection from photo-damage and viral-bacterial attack further explain their high diversity and broad spectrum ecology. The co...