Morphology of ctenostome bryozoans: 3. Elzerina, Flustrellidra, Bockiella.

Ctenostome bryozoans are a small group of bryozoans whose soft-tissue morphology has received only little attention. The present study represents the third in a series of articles dealing with the morphology of this clade of bryozoans. The morphology of three genera of Alcyonidioidea, that is, Bockiella (Alcyonidiidae), Elzerina and Flustrellidra (both Flustrellidridae), are analyzed using histology and 3D-reconstruction techniques. The general zooidal morphology is similar and externally differs by the shape of the aperture. Zooids of Elzerina binderi are elongated in the fronto-basal axis, whereas the other two are more flattened in this axis. All species show multiple pore-complexes in their zooidal walls ranging from ~66 in E. binderi, to ~30 in F. hispida and to less than 10 in Bockiella. The aperture is bilabiate in flustrellidrids and roundish in Bockiella. Apertural muscles are present as parieto-diaphragmatic muscles. The flustrellidrids have a large frontal duplicature band that further splits into four separate bands. The collar is diaphragmatic in Bockiella, but vestibular in the flustrellidrids. Lophophores are similar among the investigated species with a rejection tract in the flustrellidrids. The digestive tract shows differences in the extent and proportions of the caecum, which is large in the flustrellidrids and small in Bockiella; the anus is vestibular in all species. A funicular muscle of variable location is present in each species. Elzerina binderi has additional thin strands emanating from the digestive tract to the body wall. The parietal muscles show a unique situation in E. binderi with five bundles being present, two laterals and one distal. Several features aid in defining characters for the entire superfamily and the families Flustrellidridae and Alcyonidiidae. Besides the shape of the aperture, the frontal duplicature band, the vestibular collar and the large caecum are important. The set of characters also confirms recent notions that Bockiella belongs to the Alcyonidiidae.