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Fly-speck fungi reproduce via thyriothecia that consist of sporogenous tissue appressed to cuticle surfaces of plant leaves and covered by a shield-like scutellum. Thyriothecial scutella likely evolved repeatedly in Dothideomycetes (Ascomycota), and their morphology varies by lineage. Fly-speck fungi have an exceptionally good fossil record that begins in the Mesozoic. The interpretation of scutellum characters in fossils may provide insights into origins of Dothideomycetes and help calibrate the timing of ascomycete evolution. From sediments of the Lower Cretaceous (125–112 Ma) Potomac Group of Virginia, from Dutch Gap Canal, lower Zone 1, we found scutella similar to those of extant Aulographaceae (Dothideomycetes), attached to a single piece of dispersed coniferous cuticle. We analyze hyphae and scutellum development among four extant Aulographaceae species for comparison with the fossil. The excellent preservation of fungi on the leaf cuticle surface allows us to infer a developmental sequence for the fossil. Scutellum development begins with coordinated growth of multiple neighboring generator hyphae and continues with hyphae producing two-dimensional pseudomonopodial, dichotomous, radial growth. Asci and ascospores were not found. We coded states for seven morphological characters using direct observations of the fossil and eight extant taxa, and using the literature for 28 others. We inferred a phylogeny using nuclear 18S and 28S rDNA of 36 extant taxa, 34 Dothideomycetes and two Arthoniomycetes. The phylogeny includes newly determined sequences from five species, two from Aulographaceae. With a branch-and-bound search, we inferred the most parsimonious placements of the fossil given the molecular tree topology. The parsimony analysis constrained by the rDNA phylogeny places the fossil taxon among stem lineages near Aulographaceae or among the known living members of Aulographaceae. We describe the fossil morphotype as Protographum luttrellii, gen. et sp. nov. The fossil provides the oldest evidence of morphological characters restricted among extant fungi to Aulographaceae. [ABSTRACT FROM AUTHOR]