The case of the diatoms and the muddled mandalas: Time to recognize diatom adaptations to stratified waters

Models used to predict future ocean ecosystem and biogeochemical behaviour depend on simplified ecological frameworks allowing the definition of plankton functional types. Foremost among such frameworks has been the mandala of Margalef. His 1978 paper has been increasingly referred to in the past decade as simplified ecological schemes have been sought to help predict the effect of climate change on phytoplankton. However, the mandala is based on an understanding of the subject that is over 40 years old, when observational studies were largely limited to the coastal ocean and to near surface waters. Furthermore, most recent reproductions of the mandala are significant oversimplifications of Margalef's original. In these simplified mandala-type constructs, diatoms, in particular, have commonly been cast as a single plankton functional type that thrive in turbulent waters and decrease in abundance with increasing stratification. On this basis, it is widely predicted that diatom productivity and hence the effectiveness of the marine biological carbon pump will decrease with climate change that is driving increased stratification of the oceans. But Margalefs original took a more refined approach and depicted diatom genera that were adapted to more stratified conditions such as those characteristic of the subtropical oligotrophic gyres. If we now draw on the vast advances in observational oceanography of recent decades it is evident that diatoms may thrive, bloom and generate significant export even in the most intensely stratified and apparently oligotrophic conditions. Indeed, some diatom species have unique adaptations to such environments. We therefore suggest that it is time to abandon oversimplified schemes and recognize the diverse ecology of diatoms.