Life in a tube: morphology of the ctenostome bryozoan Hypophorella expansa.

Bryozoa is a large phylum of colonial aquatic suspension feeders. The boring ctenostome Hypophorella expansa is unique and inhabits parchment-like polychaete tubes. Morphological studies date back to the nineteenth century, but distinct adaptations to this specific habitat have not been properly analysed, which prompted us to reexamine the morphology of this recently encountered species. The colony of H. expansa is composed of elongated stolonal kenozooids with a distal capsule-like expansion. A median transversal muscle is present in the latter, and one autozooid is laterally attached to the capsule. Unique stolonal wrinkles are embedded in the thin parts of the stolons. Single autozooids are attached in an alternating right-left succession on subsequent stolons. Polypide morphology including digestive tract, muscular system and most parts of the nervous system are similar to other ctenostomes. The most obvious apomorphic features of Hypophorella are space balloons and the gnawing apparatus. The former are two fronto-lateral spherical structures on autozooids, which provide space inside the tube. The latter perforates layers of the polychaete tube wall and consists of two rows of cuticular teeth that, together with the entire vestibular wall, are introvertable during the protrusion-retraction process. The apertural muscles are in association with this gnawing apparatus heavily modified and show bilateral symmetry. Adaptations to the unique lifestyle of this species are thus evident in stolonal wrinkles, autozooidal space balloons and the gnawing apparatus. The growth pattern of the colony of H. expansa may aid in rapid colonization of the polychaete tube layers.