Adaptations of araphid pennate diatoms to a planktonic existence

Pennate diatoms are monophyletic. Their principal cell wall elements, called valves, are shaped like a ship's hull. Within the pennates, the araphids are paraphyletic; they possess rimoportulae and pore fields located at the valve apices. The pore fields exude mucilage pads with which cells attach to one another to form chains. Many taxa use the pads also for attachment to substrata. Only a few genera are truly planktonic. The main question addressed in this study is whether the planktonic lifestyle is ancestral or derived. Phylogenies inferred from nuclear SSU rDNA gene sequences of diatoms indicated that the attached lifestyle is ancestral among the araphids, whereas a typically planktonic lifestyle seems to have developed at least three times and possibly four times independently. Acquisition of a planktonic lifestyle from benthic ancestry was accompanied by a reduction in the silicification of cell-wall elements, but changes in morphological characters shared by all four clades were not detected. The reason why only three or four araphid pennate clades have adopted a planktonic lifestyle may be related to constraints associated to their sexual reproduction mode. Partner cells of opposite mating type align with one another and produce isogametes. These gametes lack flagella; they move to one another in an amoeboid fashion, which functions well on surfaces, but seems a liability in a turbulent water column. The planktonic lineages must have overcome this constraint, e.g. by sinking to the bottom, or aggregating, to perform sexual reproduction. Members of the four araphid pennate lineages are now common constituents of the plankton, suggesting that they are ecologically successful.