Ultrastructure of the osphradium of the Tertiary relict snail, Campanile symbolicum Iredale (Mollusca, Streptoneura)

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Abstract
The osphradium of Campanile symbolicum Iredale, 1917 is a gill-like, bipectinate sense organ, which is located at the left side of the mantle roof. The mass of the deeply clefted sensory epithelium of the leaflets is built up by sensory cells, which are provided with deeply invaginated aberrant cilia and large cytosomes containing pigment formations. In addition, many free nerve processes are present, bearing a single or few sensory cilia with accessory centrioles. Polyciliary cells are interspersed. A cell type with netlike or concentrically arranged smooth endoplasmic reticulum is commonly found near the central axis of the osphradium. The central zone of each leaflet includes nervous tissue and a complicated muscular grid, with pore cells and fibroblasts also present. Based on the fine-structural data the functional and ecological significance of the osphradium of Campanile symbolicum is discussed. The combination of herbivory and a lamellar osphradium is rare among the Gastropoda, suggesting that the osphradium of Campanile might also be involved in reproductive biology. Many fine-structural features of the osphradium of Campanile symbolicum are unique among the gastropods and reflect the phylogenetic isolation of this relict snail. The net-like cell type, however, is probably homologous with the so-called Si4 cell in the remaining caenogastropods, for which a largely different osphradial fine-structure is diagnostic. The affinities of Campanile symbolicum are probably closer to the Caenogastropoda than to the Allogastropoda and Euthyneura. With present knowledge it might be best classified near the base or even as the first clade within the Caenogastropoda.