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Smittium, one of the first described genera of gut fungi, is part of a larger group of endosymbiotic microorganisms (Harpellales) that live predominantly in the digestive tracts of aquatic insects. As a diverse and species-rich taxon, Smittium has helped to advance our understanding of the gut fungi, in part due to the relative success of attempts to culture species of Smittium as compared to other members of Harpellales. Approximately 40% of the 81 known species of Smittium have been cultured. This is the first Smittium multigene dataset and phylogenetic analysis, using the 18S and 28S rRNA genes, as well as RPB1, RPB2, and MCM7 translated protein sequences. Several well-supported clades were recovered within Smittium. One includes the epitype S. mucronatum (the “True Smittium” clade), and another contains many species including S. simulii and S. orthocladii (the “Parasmittium” clade). Ancestral states …