Re-discovery and novel contributions to morphology and multigene phylogeny of Myxophyllum steenstrupi (Ciliophora: Pleuronematida), an obligate symbiont of terrestrial pulmonates

Abstract

Abstract Myxophyllum steenstrupi is a symbiotic ciliate living in the body slime and mantle cavity of terrestrial pulmonates (Gastropoda: Pulmonata). In the present study, M. steenstrupi was re-discovered after almost 30 years and characterized using an integrative morpho-molecular approach for the first time. Myxophyllum is distinguished by a broadly ovate, about 140 × 115 μm-sized body, a nuclear apparatus typically composed of seven macronuclear nodules and a single micronucleus, a central contractile vacuole, a shallow oral cavity situated in the posterior body region and dense somatic ciliature with extensive thigmotactic field. According to the present phylogenetic analyses of two mitochondrial and five nuclear markers, M. steenstrupi is classified in the predominantly free-living order Pleuronematida (Oligohymenophorea: Scuticociliatia). This order also encompasses other taxa isolated from molluscs and traditionally classified along with Myxophyllum in the order Thigmotrichida. The proper classifications of Myxophyllum was hampered by the dramatic remodelling of its oral apparatus (reduction of the paroral membrane and adoral organelles, formation of vestibular kineties), a transformation that was likely induced by its firm association with terrestrial gastropods. The present study also documents that various ciliate lineages independently became commensals or parasites of various aquatic and terrestrial molluscs.