North American Paleozoic land snails, with a summary of other Paleozoic nonmarine snails

Abstract

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. The apertural barriers in Anthracopupa are identical in placement and growth pattern with those of living Tornatellinidae and independently confirm the family placement derived from study of the general form. One new species, A. sturgeoni, has been named. Anthracopupa is found most commonly in thin limestones interpreted as having been deposited in pools into which the small shells floated. Dendropupa is most commonly found in erect tree stumps that were covered by rapid sedimentation. Both environments are similar to those in which the shells of allied living species may be found today, and the fossils support environmental interpretations made entirely from lithology. A survey of the few European occurrences of Paleozoic land snails indicates that both Anthracopupa and Dendropupa occur in Lower Permian strata; Anthracopupa is known from beds as old as Westphalian B. These genera cannot be used for determining the Carboniferous-Permian boundary. Both the long local stratigraphic range of A. brittanica and D. vetusta reported in the literature and the moderately long range and great variability of A. ohioensis suggest that the land snails have little stratigraphic utility. On the other hand, the occurrence of these land snails in the late Paleozoic of the Northern Hemisphere provides further fossil evidence suggestive of a closed Atlantic Ocean at that time. A comparison of the Paleozoic and the present distributions of land -snail families on both sides of the Atlantic provides some interesting data on geographic shifts of organisms. Finally, the assignment of the earliest land snails to extant taxa at the family level indicates that the subclass Pulmonata has been very conservative in its evolution after initial radiation. A few notes on Paleozoic freshwater snails complete this survey.