Micah Dunthorn

Ciliates have a long history of being central in evolutionary and ecological studies on eukaryotic microorganisms. Although thousands of species have been discovered, their total diversity still remains unknown. Here, we will discuss two unsolved problems that hinder the further exploration of ciliate diversity at the species level, and potential solutions to these problems are proposed. First, ciliate morphospecies are difficult to identify because the different silver stains are not scalable (they do not represent high-throughput methods) and basic supplies are lacking (e.g., protargol); a solution may be the development of fluorescent staining techniques. Second, ciliate phylogenetic species are difficult to identify because of extensive paralogy in nuclear-protein-coding genes; a solution may be to concentrate on sequencing mitochondrial genomes. These two approaches could be integrated into a high-throughput fluorescent-single-cell sorting and mitochondrial genomes sequencing process that would enable the observation and better understanding of ciliate species on a massive scale.

Mitochondrial SSU-rDNA has been used recently to infer phylogenetic relationships among a few ciliates. Here, this locus is compared with nuclear SSU-rDNA for uncovering the deepest nodes in the ciliate tree of life using broad taxon sampling. Nuclear and mitochondrial SSU-rDNA reveal the same relationships for nodes well-supported in previously-published nuclear SSU-rDNA studies, although support for many nodes in the mitochondrial SSU-rDNA tree are low. Mitochondrial SSU-rDNA infers a monophyletic Colpodea with high node support only from Bayesian inference, and in the concatenated tree (nuclear plus mitochondrial SSU-rDNA) monophyly of the Colpodea is supported with moderate to high node support from maximum likelihood and Bayesian inference. In the monophyletic Phyllopharyngea, the Suctoria is inferred to be sister to the Cyrtophora in the mitochondrial, nuclear, and concatenated SSU-rDNA trees with moderate to high node support from maximum likelihood and Bayesian inference. Together these data point to the power of adding mitochondrial SSU-rDNA as a standard locus for ciliate molecular phylogenetic inferences.

Three putative ciliate fossils were described from the Neoproterozoic Doushantuo Formation in China: Eotintinnopsis, Wujiangella, and Yonyangella. The identity of these fossils is important for our understanding of the origins and early morphological evolution within ciliate clades. Here we compare the homology of the fossil characteristics with those in their proposed ciliate relatives. Eotintinnopsis resembles a tintinnid, but its feathery tentacle-like apical structure is probably not homologous within any known ciliate. Wujiangella presents homology issues with the size and distribution of its putative somatic cilia. Yonyangella appears to be a suctorian with its tentaclelike structures, but the presence and size of its putative somatic cilia pose homology issues. We suggest that these three fossils are likely to be taphonomically and diagenetically distorted and altered acritarchs. These alterations include secondary mineral encrustations on the interiors of vesicles, the crushing, folding and other distortions of the vesicles, the bending and crushing of the acritarch spines, and the preservation of organic material in and outside of the cysts. The earliest known ciliate fossil remains a tintinnid that occurs in the Ordovician of Kazakhstan.