Morpho-molecular characterization of the litostomatean predatory ciliate Phialina pupula (Müller, 1773) foissner, 1983 (haptoria, lacrymariidae)

The morphology and phylogenetic position of a haptorian ciliate, Phialina pupula (Müller, 1773) Foissner, 1983, isolated from microaerobic sandy sediments of the floodplain area of the Boise River, Idaho, U.S.A., were studied using live observation, protargol impregnation, scanning electron microscopy, and the 18S rRNA gene as well as the ITS region. The Boise population of P. pupula is characterized by a size of about 60–130 × 20–50 µm, an elliptical macronucleus with a single micronucleus, highly refractive dumbbell-shaped inclusions scattered throughout the cytoplasm and concentrated in the anterior body half, a single subterminal/terminal contractile vacuole, about 10 µm long rod-shaped extrusomes, and an average of 15 ciliary rows. In phylogenetic analyses, the newly obtained sequences from P. pupula and Lacrymaria olor clustered within the family Lacrymariidae with full to moderate statistical support. Neither the genus Phialina nor the genus Lacrymaria was depicted monophyletic both in the single gene and multigene phylogenetic inferences. Specifically, the genus Phialina was shown as a paraphyletic assemblage containing members of the polyphyletic genus Lacrymaria. This indicates that the phialinid bauplan, i.e., an anterior body end differentiated into a head-like structure directly attached to the trunk, might represent the ground pattern in the family Lacrymariidae. On the other hand, the long highly contractile neck carrying the head-like structure probably evolved later and convergently in multiple Lacrymaria species from Phialina-like ancestors.