Oldest fossil ciliates from the Cryogenian glacial interlude reinterpreted as possible red algal spores

The Cryogenian Period experienced two long lived global glaciations known as Snowball Earths. While these events were dramatic, eukaryotic life persisted through them, and fossil evidence shows that eukaryotes thrived during the ca. 30-million-year interlude between the glaciations. Carbonate successions have become an important taphonomic window for this interval. One of the most notable examples is the ca. 662–635 Ma Taishir Formation (Tsagaan Olom Group, Zavkhan Terrane, Mongolia) which has yielded a number of eukaryotic fossil taxa. Here, we examine more closely the morphology and taxonomic affinity of some of these Taishir fossils previously interpreted as remains of ciliate tintinnid loricae (purportedly the oldest fossil ciliates). New morphological and ultrastructural analyses indicate that these fossils are not ciliate tintinnids. Instead, we propose a new interpretation: that they are algal reproductive structures related to coeval macroscopic organic warty sheets described as...