Zone-diagnostic Middle to Upper Devonian ammonoids and conodonts occur in a sequence of thin carbonate beds in siliciclastic sections along the Allegheny front and northern fold belt of central Pennsylvania. The lowest, in the upper Tully Formation, and the succeeding four in the Harrell Shale, are southward continuations of marker beds long known in the Tully Limestone and overlying basin, slope, and shelf facies of the Genesee Group of New York State. These pelagic beds are Appalachian Basin signals of transgressive condensed intervals within global high-sea-level episodes and are associated with a worldwide biotic overturn in the tropical marine realm.

The Upper Givetian Tully Pharciceras amplexum bed has close lithic and faunal equivalents in the Taghanic biocrisis interval in Morocco and southern Europe. The Lower Frasnian styliolinid upper Genundewa Limestone, with the entry of Manticoceras, and, in the succeeding dark West River Shale, the Bluff Point Bed, with the entry of late Koenenites and early palmatolepid conodonts, are Appalachian highstand signals of the worldwide Timan bioevent transgression recognized in Western Australia, the Russian Platform, southern Europe, Morocco, western Canada, and now Pennsylvania. The widespread occurrences of discrete ammonoid beds, even in the more proximal or shoreward sedimentary settings of the Catskill Delta, reinforce the view that they accumulated under conditions of sequestered sediment supply when transgressions flooded the newly vegetated and deep-root-forested delta flats. As such, they overprinted or interrupted the westward progradation of siliciclastic sediments generated by mountain-building tectophases of the Acadian orogeny.

The Lower Frasnian ammonoid Epitornoceras dennisoni n. sp., described herein, from the Crosby Beds in the Harrell Shale at Milesburg, Pennsylvania, is named in honor of John Dennison.