Relationships of Diatoms to Other Groups
Despite a number of studies to examine phylogeny, using one or several genes, the relationships of diatoms to other groups are still unclear and there is still a huge gap in our understanding of how and when diatoms acquired their unusual morphology and life-cycle characteristics. The diatoms have often been treated as a separate phylum, reflecting their unique features. Pascher (1914, 1921) suggested that the diatoms have features in common with the Chrysophyceae and Xanthophyceae and therefore placed these classes and the Bacillariophyceae in the phylum Chrysophyta. Ultrastructural and molecular sequence data have confirmed the general thrust of Pascher’s idea, placing the diatoms unambiguously among the heterokont protists (‘stramenopiles’) within the chromalveolates (Adl et al. 2005).
In the past, it was sometimes suggested that diatoms evolved well before their appearance in the fossil record and that the early phases in diatom evolution were lost long ago through diagenesis of diatomites to chert (e.g. Round 1981). This is made extremely unlikely by recent molecular phylogenies, which date the origin of diatoms towards the beginning of the Mesozoic Era. Furthermore, a close relationship to other silica scale or silica skeleton-producing algae and protists, such as the Chrysophyceae, is not evident in recent analyses. The closest known relatives of the diatoms are the bolidophytes (Bolidophyceae), which are a small group of marine autotrophic picoplankton with the same kind of plastids and flagellum structure as diatoms and some other autotrophic heterokonts (Guillou et al. 1999). However, bolidophyte cells are highly reduced and simplified and do not seem to produce any silica structures, although it is possible that silicifying life cycle stages have been missed.
Mann and Marchant (1989) suggested that another group, the Parmophyceae, may also be closely related to diatoms and thus may give hints as to how diatoms arose, because they produce silica scales that in some respects (radial pattern subtended by a central ring, space-filling development of pattern) resemble diatom valves and girdle bands. So far, no DNA sequences have been confirmed to be derived from Parmophyceae, but a clade of unknown heterokonts closely related to diatoms and bolidophytes has been detected by Lovejoy et al. (2006) and may represent the Parmophyceae; it is certainly important for understanding the evolution of both bolidophytes and diatoms that the organisms detected by Lovejoy et al. are fully characterized.
Round and Crawford (1981) and Mann and Marchant (1989) developed hypotheses about how the diatom frustule evolved, based on comparative morphology. Both suggested that diatoms probably arose from scaly celled ancestors. The scale-case was thought initially to have been homogeneous (all the scales were fairly alike in size, shape and structure). Then there was a stage in which the scales became differentiated into larger valve-like scales and narrower ones that resembled the segments found in the girdles of modern Rhizosolenia species (though this is not meant to imply that modern rhizosolenids are a basal offshoot), and a still later stage when the proto-girdle bands became even narrower, forming hoops around the cell.
According to this evolutionary progression, valves and girdle bands would have a common origin, which seems reasonable because their structure is often similar and they are formed in similar ways. Furthermore, cells covered evenly with scales are known in diatoms, in the auxospores of some centric diatoms, e.g. Melosira or Ellerbeckia (Crawford 1974, Schmid & Crawford 2001).
The main differences between the Round–Crawford and Mann–Marchant hypotheses are in the assumptions made about the nature of the scales and scaly cell in the early (‘Ur’) diatoms and the nature of the scaly cells themselves. In the Mann–Marchant scheme, the scales of the pre-diatom were space-filling structures, which abutted to form the complete, functional cell wall of a temporarily dormant cyst, whereas Round and Crawford envisaged the scales as separate elements that did not abut but were imbricate, covering growing vegetative cells as in modern synurophytes.
No precursors of diatoms have been identified from the fossil record.