Interactive 3D anatomy and affinities of the Hyalogyrinidae, basal Heterobranchia (Gastropoda) with a rhipidoglossate radula

Whereas Hyalogyrina Marshall, 1988 was originally considered a skeneid vetigastropod, the family Hyalogyrinidae War,n & Bouchet, 1993 has later been classified as basal Heterobranchia despite their rhipidoglossate radula. In order to evaluate this placement and to shed more light on the origin of all higher Gastropoda, we investigated five representatives of all three nominal hyalogyrinid genera by means of semithin serial sectioning and computer-aided 3D reconstruction of the respective anatomy, which we present in an interactive way. In general the morphological features (shell, external morphology, anatomy) fully confirm the placement of Hyalogyrinidae in the Heterobranchia, but in particular the conditions of the genital system vary substantially within the family. The ectobranch gill of Hyalogyrinidae is shared with Valvatidae, Cornirostridae, and Xylodisculidae; consequently all these families are united in Ectobranchia Fischer, 1884. The rhipidoglossate hyalogyrinid radula suggests independent acquisition of taenioglossate radulae in the Caenogastropoda and other Ectobranchia. Therefore, the origin of the Heterobranchia-and thus of all higher gastropods-looks to have taken place already on the rhipidoglossate, i.e. the 'archaeogastropod', level of evolution. Ectobranchia are considered the first extant offshoot of the Heterobranchia; implications for the stem species of the latter are outlined.