Evolution of the diatoms: major steps in their evolution and a review of the supporting molecular and morphological evidence

Abstract:

Over the years, many reviews of different aspects of diatom biology, ecology and evolution have appeared. Since 1993 many molecular trees have been produced to infer diatom phylogeny. In 2004, Medlin & Kaczmarska revised the systematics of the diatoms based on more than 20 years of consistent recovery of two major clades of diatoms that did not correspond to a traditional concept of centrics and pennates and established three classes of diatoms: Clade 1 = Coscinodiscophyceae (radial centrics) and Clade 2 = Mediophyceae (polar centrics + radial Thalassiosirales) and Bacillariophyceae (pennates). However, under certain analytical conditions, an alternative view of diatom evolution, a grades of clades, has been recovered that suggests a gradual evolution from centric to pennate symmetry. These two schemes of diatom evolution are evaluated in terms of whether or not the criteria advocated by Medlin & Kaczmarska that should be met to recover monophyletic classes have been used. The monophyly of the three diatom classes can only be achieved if (1) a secondary structure of the small subunit (SSU) rRNA gene was used to construct the alignment and not an alignment based on primary structure and (2) multiple outgroups were used. These requirements have not been met in each study of diatom evolution; hence, the grade of clades, which is useful in reconstructing the sequence of evolution, is not useful for accepting the new classification of the diatoms. Evidence for how these two factors affect the recovery of the three monophyletic classes is reviewed here. The three classes have been defined by clear morphological differences primarily based on gametangia and auxospore ontogeny and envelope structure, the presence or absence of a structure (tube process or sternum) associated with the annulus and the location of the cribrum in those genera with loculate areolae. New evidence supporting the three clades is reviewed. Other features of the cell are examined to determine whether they can also be used to support the monophyly of the three classes.

Key Words::

• Auxospores
• Diatoms
• Evolution
• Frustules
• Phylogeny