The unplumatellid Plumatella fruticosa found its home: Hirosella gen. nov. morphological arguments for the systematic placement of a freshwater bryozoan

Abstract

Bryozoans are colonial, suspension‐feeding lophotrochozoans. The phylum consists of the large group of chiefly marine Myolaemata and the exclusively limnic Phylactolaemata. Each colony consists of individual zooids that comprise the protective cystid and the retractable polypide. Phylactolaemates are a small group of approximately 90 species in 6 families. They feature a body wall, that can either be gelatinous, as in the families Stephanellidae, Lophopodidae, Cristatellidae and Pectinatellidae, or encrusted, as in Plumatellidae and Fredericellidae. Morphological investigations of the most specious plumatellids are rare and focus on few species. Plumatella fruticosa is of particular interest in this regard, as it shows a mosaic of plumatellid and fredericellids characters. The most recent phylogeny clusters P. fruticosa with cristatellids and pectinatellids as sister groups to fredericellids. Hence, there is considerable doubt, whether P. fruticosa is truly a plumatellid. Therefore, this study aims to reinvestigate the morphology of P. fruticosa with confocal microscopy and section‐based three‐dimensional reconstruction. The new data show that P. fruticosa has numerous conspicuous stumps from fragmented proliferation buds, which are otherwise only known from fredericellids. Like fredericellids, P. fruticosa grows erect, but in contrast, has a horseshoe‐shaped lophophore and floatoblasts. Besides the proportions of the lophophore, the tentacle sheath and digestive tract resemble a fredericellid‐like situation. Myoanatomical details like the pronounced longitudinal muscles of the vestibular wall and tentacle sheath differ from plumatellids and favour the recently proposed scenario, which places P. fruticosa next to Pectinatellidae and Cristatellidae. In addition, the intertentacular membrane of P. fruticosa shows structural similarity to cristatellids as it is attached to the tentacles via lamellae. Taking all aspects into account, we erect a new family: Hirosellidae fam. nov. including the new genus Hirosella gen. nov.