North American Paleozoic Land Snails with a Summary of Other Paleozoic Nonmarine Snails

Land snails from the Paleozoic of North America are known from the coal fields of eastern Canada, from the Dunkard basin west of the Allegheny Mountains, and from the western margin of the Illinois basin. The earliest finds were made about 125 years ago; essentially no new information has been recorded for a century. Large collections of Anthracopupa from the Dunkard basin sparked inquiry into the land snails from the other two areas. Studies using the SEM (scanning electron microscope) have provided considerable insight into microdetails of shell structure, which allow systematic assignment of these gastropods. All may be assigned to extant families, except one, for which insufficient material allows only superfamily assignment. The prosobranch Dawsonella is confirmed as being a terrestrial neritacean gastropod. To date, it is known only from the upper Middle Pennsylvanian of Illinois and Indiana. All the other Paleozoic land snails are stylommatophoran pulmonates; their current classification as nonmarine cyclophoraceans is not correct. Restudy of material from the Joggins section of Nova Scotia indicates that representatives of two ordinal groups of pulmonates appeared simultaneously in upper Lower Pennsylvanian strata; the oldest land prosobranch is found in only very slightly younger rocks. Zonites (Conulus) priscus is reassigned to the new genus Protodiscus in the extant family Discidae. Dendropupa is placed within the family Enidae, Anthraaopupa is placed in the family Tornatellinidae, and 'Pupa' bigsbii is assigned to the superfamily Pupillacea. All four of these family-level taxa are diverse and belong to two orders within the superorder Stylommatophora, heretofore considered a derived rather than an ancestral stock. Anthracopupa ohioensis Whitfield is a highly variable species, and two other species Naticopsis (?) diminuta and A.(?) dunkardona, both named by Stauffer and Schroyer, are placed in synonymy with it. To obtain taxonomic data to support the family placement of Anthracopupa, growth forms of modern pupillid and tornatellinid snails have been distinguished.