Foraminiferal wall microstructures, consistent with the molecular-based high-rank classification, are critical to understanding foraminiferal evolution and advanced taxonomic relationships. Although test structures are well documented for recent, Cenozoic, and some Mesozoic foraminifera, the diagnostic characteristics of Paleozoic taxa are largely unexplored. The majority of calcareous Paleozoic foraminifera have been assigned to the Fusulinata based on questionable homogeneously "microgranular" test wall microstructures, which have never been sufficiently documented for most taxa. We investigated the test structures of exceptionally well-preserved Devonian (Eifelian) Semitextularia thomasi, representing the first calcareous true multichambered (serial) foraminifera, and compared this species with a large fusiform Permian representative of "true" fusulinids (Neoschwagerinidae). The tests of Semitextularia thomasi display lamellar structures that are not observed in any other fossil or recent foraminiferal group. The Paleozoic foraminifera, traditionally referred to one taxon (the class Fusulinata), possess at least three contrasting test wall microstructures, representing separate high-rank taxonomic groups. Fusulinata is most likely a highly polyphyletic group that is in need of taxonomic revision. The term Fusulinata, defined as including all Paleozoic calcareous forms except Miliolida and Lagenata, is not phylogenetically meaningful and should no longer be used or should be restricted to true complex fusulinids with microgranular test structures, which appeared in the Carboniferous.
Keywords: biomaterial; foraminifera; fusulinids biomineralization; nanostructures.