Anatomy and ultrastructure of ventral pharyngeal organs and their phylogenetic importance in Polychaeta (Annelida)

Summary A comparative anatomical and ultrastructural study of ventral pharyngeal organs (pharyngeal bulbs) was carried out in two species of the Dinophilidae: Dinophilus gyrociliatus and Trilobodrilus axi. Special attention was paid to the fine structure of the stomodeal epithelium, cuticle, glands, muscles, and myoepithelial junctions. The differences between the species are very slight. The pharyngeal organ of the Dinophilidae is characterized by the following features: solid muscle bulbus made up of muscle cells only, bulbus muscle cells with two myofilament systems crossing at an angle of about 90°, gap junctions between these muscle cells, bulbus projects into a pharyngeal sac and bears rostrally a specific epithelium and cuticle, no bulbus glands, no investing (= sagittal) muscles, specific cuticle ultrastructure, cilia of ascending oesophagus with asymmetric tips, specific structure and position of salivary gland openings. The phylogenetic importance of these structures is discussed. Some of these characters are clearly autapomorphic features of the Dinophilidae and no common derived structures to other families with a ventral pharyngeal organ are present. Therefore, it is most likely that the dinophilid pharyngeal organ evolved independently. These findings do not agree with the hypothesis of the unity of the archiannelid families (Polygordiidae, Protodrilidae, Saccocirridae, Nerillidae, Dinophilidae, and Diurodrilidae) established on the basis of an assumed structural similarity of their ventral pharyngeal organs.Abbreviations bb basal body - bep bulbus epithelium - bl basal lamina - bm bulbus muscle - c cilium - cc coelenchyme cell - cm circular muscle - cr caudal rootlet - cu cuticle - dblm dorsal bulbus longitudinal muscle - dlm dorsal longitudinal muscle - dsn dorsal stomatogastric nerve - dy dyad - el electron-dense layer - fl fibrous layer - fi filaments - g Golgi apparatus - gl gland cell - hv homogeneous vesicle - l lipid droplet - la external lamina - lal lamellar layer - ll lower lip - lm longitudinal muscle - ly lysosome - m mitochondrion - mo mouth opening - mt microtubule - mv microvillus - mvp microvillar process - n nucleus - nu nucleolus - oes oesophagus - pcom preoral commissure - phf pharyngeal fold - phl pharyngeal lumen - phs pharyngeal sac - pms peripheral myofilament system - r rootletlike structure - rer rough endoplasmic reticulum - rr rostral rootlet - s sarcoplasmic reticulum - sc salivary canal - scom suboesophageal commissure - sd septate desmosome - ser smooth endoplasmic reticulum - sg secretory granule - sgl salivary gland - sn stomatogastric nerve - st stomach - step stomodeal epithelium - tep transitional epithelium - tf tonofilaments - va vacuole - vlm ventral longitudinal muscle - vsn ventral stomatogastric nerve - z z-element - za zonula adherens