Reconstruction of Evolutionary History of Pleurostomatid Ciliates (Ciliophora, Litostomatea, Haptoria): Interplay of Morphology and Molecules

Abstract

Pleurostomatids are raptorial ciliates that form a very distinct group within the Haptoria. Traditionally, the order Pleurostoma- tida was divided into two families: the Amphileptidae with two perioral kineties and a suture formed by the right side ciliary rows, and the Litonotidae with three perioral kineties and without suture. However, molecular phylogenies depicted the traditional Amphileptidae as a paraphyletic assemblage nesting also the Litonotidae. To overcome this problem we have analyzed genealogy of pleurostomatids using morphological data and 18S rRNA gene sequences, including newly sequenced genera Acineria and Kentrophyllum. Specifically, we have combined a morphological and molecular approach and have used also some other phylogenetic tools such as phylogenetic networks, split spectrum analysis, quartet mapping as well as the likelihood method of tracing history of morphological characters. These analyses show that: (1) there are not two but three distinct pleurostomatid lineages - Epiphyllidae fam. nov., Amphileptidae and Litonotidae; (2) epiphyllids (Epiphyllum + Kentrophyllum) represent a basal pleurostomatid group which is defined by two perioral kineties, by the presence of a suture on both the right and the left side of the body, by the loss of the oral bulge extrusomes, and by the extrusome fringe extending all around the body except for the oral region; (3) the families Amphileptidae and Litonotidae are monophyletic each, and represent sister groups; (4) Acineria belongs to the Litonotidae, as already indicated by morphological data; (5) Loxophyllum is a monophyletic and crown genus of the Litonotidae; and (6) Litonotus is paraphyletic, which could be very likely caused by a rapid radiation event that did not allow primary nucleotide homologies to be fixed.