Oral Monokinetids in the Free-Living Haptorid Ciliate Enchelydium polynucleatum (Ciliophora,Enchelyidae): Ultrastructural Evidence and Phylogenetic Implications1

Special ultrastructural characteristics of the haptorid soil ciliate Enchelydium polynucleatum Foissner, 1984 are the restriction of the parasomal sacs to the area of the “brush” and finger-like projections of the food vacuole membrane into the lumen of the vacuole. The general organization of the infraciliature is similar to that of Spathidium and some buetschliids because the anterior ends of the somatic kineties are condensed and obliquely bent. Enchelydium is similar to haptorids and buetschliids in possessing monokinetid somatic fibrillar structures with the classical fibrillar associates: 1) a short kinetodesmal fiber; 2) two transverse microtubular ribbons; 3) a long postciliary microtubular ribbon; and 4) a system of overlapping subkinetal microtubules, which seems to be absent in the buetschliids. Unlike Spathidium and all other haptorids so far investigated ultrastructurally, serial sections show that there are no oral dikinetids, as in the endocommensal buetschliids and balantidiids. Instead, three to six anterior kinetids in each ciliary row have nematodesmal bundles extending into the cytoplasm and surrounding the cytopharynx. These kinetids lack cilia and all fibrillar associates except enlarged transverse ribbons, which extend anteriorly and inwards to support the cytopharynx. Other similarities between the buetschliids and Enchelydium are the conspicuous rough endoplasmic reticulum and abundant sausage-like vesicles in the oral region. As in other haptorids, Enchelydium has two types of toxicysts and one type of mucocyst. These observations strongly suggest that Enchelydium belongs to the ancestral stock of both the Haptorida and the Archistomatida. The similarities in the somatic and oral infraciliature and ultrastructure of the Haptorida and the Archistomatida suggest that they belong to the same subclass, Haptoria Corliss, 1974.