Ciliates — Protists with complex morphologies and ambiguous early fossil record

Since ciliates rarely possess structures that easily fossilize, we are limited in our ability to use paleontological studies to reconstruct the early evolution of this large and ecologically important clade of protists. Tintinnids, a group of loricate (house-forming) planktonic ciliates, are the only group that has a significant fossil record. Putative tintinnid fossils from rocks older than Jurassic, however, possess few to no characters that can be found in extant ciliates; these fossils are best described as ‘incertae sedis eukaryotes’. Here, we review the Devonian fossil Nassacysta reticulata and propose that it is likewise another incertae sedis eukaryote due to the lack of any unambiguous ciliate characters. Future tintinnid fossil descriptions would be most helpful if: (i) neutral terminology is used in the descriptions but ciliate-specific terminology in the interpretations; (ii) the current ciliate classification is used, although fossil data may expand or modify classifications based on modern forms; (iii) close collaboration with specialists studying extant ciliates is done; and (iv) editors include an expert of extant ciliates in the review process.