North American Paleozoic 

Land Snails, With a Summary of 

Other Paleozoic Nonmarine

GEOLOGICAL SURVEY PROFESSIONAL PAPER 1072
North American Paleozoic 

Land Snails, With a Summary of 

Other Paleozoic Nonmarine Snails

By ALAN SOLEM and ELLIS L. YOCHELSON

GEOLOGICAL SURVEY PROFESSIONAL PAPER 1072

A redescription and review of Pennsyluanian 

and Early Permian terrestrial forms, 

demonstrating the early diversification 

of the Pulmonata

UNITED STATES GOVERNMENT PRINTING OFFICE, WASHINGTON: 1979
UNITED STATES DEPARTMENT OF THE INTERIOR 

CECIL D. ANDRUS, Secretary

GEOLOGICAL SURVEY 

H. William Menard, Director

Library of Congress Cataloging in Publication Data

Solem, George Alan, 1931-

North American Paleozoic land snails.

(Geological Survey professional paper 1072)

Bibliography: p. 36-38.

Includes index.

1. Gasteropoda, Fossil. 2. Paleontology Paleozoic. 3. PaleontologyNorth America. I. Yochelson, Ellis Leon, 1928- joint author. II. Title. 

III. Series: United States. Geological Survey. Professional paper ; 1072.

QE808.S67 564'.3'097 78-606004

For sale by the Superintendent of Documents, U.S. Government Printing Office

Washington, D.C. 20402

Stock Number 024-001-03197-3
CONTENTS

Page

Abstract. ______________________________ 1

Introduction ____________________________ 1

Acknowledgments ______________________ 2

Earlier systematic work _____________________ 3

Stratigraphic setting and local occurrence ___________ 3

Illinois and Indiana _____________________ 3

Ohio, Pennsylvania, and West Virginia __________ 5

New Brunswick and Nova Scotia _____________ 6

Analysis and classification of taxa _______________ 7

Other Paleozoic nonmarine snails _______________ 26

Systematic summary ______________________ 28

Phyletic significance _______________________ 29

Ecological significance ______________________ 31

Biogeographic significance ___________________ 33

Register of localities for Monongahela-Dunkard

gastropods ___________________________ 34

References cited _________________________ 36

Index _______________________________ 39

ILLUSTRATIONS

[Plates follow index]

PLATE 1. Strophella, Anthracopupa, and ?'Anthracopupa.

2. Protodiscus and Pupa.

3. Anguispira.

4. Dendropupa.

5-7. Anthracopupa.

8. Protodiscus, Anthracopupa, and Dendropupa.

9. Anthracopupa.

10. Dawsonella.

Page

FIGURE 1. Shell structures in Dendropupa and a modern enid ______________________________________ 11

2. Original illustrations of Dendropupa primaeva _________________________________________ 14

3. Polished section of Anthracopupa ohioensis ___________________________________________ 18

4. Range of variation in barrier development of Anthracopupa ohioensis ____________________________ 19

5. Ontogenetic development of expanded lip and palatal ridges in Anthracopupa ohioensis _________________ 20

6. Shell-form differences in the pupillid and tornatellinid taxa _________________________________ 22

TABLES

Page

TABLE 1. General correlation of parts of the Pennsylvanian and Permian between North America and Europe, showing the approximate positions of the stratigraphic groups from which land snails have been collected in North America. _ _ 4

2. Size (in mm) and shape variation in samples of Anthracopupa ohioensis Whitfield, 1881 _________________ 16

3. Frequency distribution for shell height, shell diameter, height/diameter ratio, and whorl count in pooled samples of adult

materials of Anthracopupa ohioensis Whitfield, 1881 ___________________________________ 17

4. Relation of shell height to shell diameter in all adult specimens of Anthracopupa ohioensis Whitfield, 1881 ______ 17

5. Relation of shell height to whorl count in all adult specimens of Anthracopupa ohioensis Whitfield, 1881 _______ 17

6. Systematic list of the Paleozoic land snails ____________________________________________ 29

III
NORTH AMERICAN PALEOZOIC LAND SNAILS, 

WITH A SUMMARY OF OTHER PALEOZOIC NONMARINE SNAILS

By ALAN SOLEM1 and ELLIS L. YOCHELSON

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, Anthracopupa 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

'Department of Zoology, Field Museum of Natural History, Chicago, 111. 60605.

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.

INTRODUCTION

The restudy of type specimens, analysis of significant new collections of Anthracopupa from Ohio, and 

examination of diverse materials from other areas of 

the world have permitted us to prepare this detailed 

revision of the North American Paleozoic land snails 

and to summarize the probable affinities, stratigraphic significance, and biogeographic implications of 

all the Paleozoic nonmarine snails. SEM (scanning 

electron miscroscope) photographs of several taxa 

have revealed details of structure in the fossils that 

permitted assignment of the Paleozoic species to extant family groupings. One plate (pi. 1) documents 

types hitherto illustrated by drawings and supplements the plates of SEM photographs that illustrate 

shell structures.

Representatives of three orders and five families of 

land snails appear essentially simultaneously in the 

fossil record of the Pennsylvanian. These families persist today, thus demonstrating that in the early late 

Carboniferous, the land snails had achieved a high 

and stable level of diversity. Only three additional orders of land snails are recognized today. One of these, 

the Systellommatophora, contains only shell-less 

slugs and has no fossil record. Land snails belonging to 

the other two orders, Mesurethra and Holopoda, do 

not appear in the fossil record until significantly later.
NORTH AMERICAN PALEOZOIC LAND SNAILS, SUMMARY OF OTHER PALEOZOIC NONMARINE SNAILS

The record of North American Paleozoic land snails 

is spotty. Several taxa (Dawsonella meeki (Bradley, 

1872), "Pupa" bigsbii Dawson, 1880, and Protodiscus 

priscus (Carpenter, 1867)) are known either from a 

single locality or a limited time span. Other taxa belong to genera (Dendropupa and Anthracopupa) that 

range from Middle Pennsylvanian (Westphalian B) 

through the Early Permian, about 45-50 million years, 

and occur in both North America and Europe.

For several taxa, the original age assignments are no 

longer accepted. We have attempted to equate the age 

of type localities with present nomenclature. As currently known, the fossil genera cannot be used to determine stratigraphic boundaries. The few species 

that are abundant have long stratigraphic ranges. Collectively, the Paleozoic land snails do provide added 

evidence that Europe and North America were joined 

as a single landmass during this time.

We were able to examine one or more specimens of 

each of the previously described North American species, except Pupa primaeva Matthew (1895), which is 

known from only one specimen that we could not locate. Study of extensive new collections from the 

Monongahela and Dunkard Groups (Upper Pennsylvanian and Lower Permian) resulted in several 

changes in synonymy and in description of a new species, Anthracopupa sturgeoni. Less intensive study of 

European and South American material has enabled 

us to make additional stratigraphic and biogeographic 

comments.

Our method of analysis of the fossil shells has been 

first to interpret, so far as possible, the basic growth 

patterns, structures, and ontogenetic developmental 

sequences. We have then compared these patterns 

and structures with the comparable ones found in extant land-snail taxa. Many of the fossils are very well 

preserved. With the aid of the SEM, we could study 

extremely fine details of the shell. The availability of 

both juvenile and adult examples permitted the determination of some aspects of shell ontogeny. When details of both structure and ontogeny in the fossils are 

found to match exactly features that are now restricted to a single extant family-level unit, we have no 

hesitation in assigning the fossil species to the extant 

family.

In the main section of this report, we describe the 

individual fossil taxa, review their structure, and suggest their individual affinities. The sequence of species is arbitrary and coincides with groupings of 

illustrations. Formal diagnoses and descriptions of 

higher taxonomic categories are included with the 

analysis of the one species in that taxon for which we 

had the greatest amount of data. This departure from 

the conventional systematic sequence of higher to

lower category is deliberate.

We then present a summary of the other Paleozoic 

nonmarine snails, which is incorporated into the sections on the phyletic, ecological, and biogeographic 

significance of our systematic conclusions.

This project is a joint one, Solem being responsible 

for the data on present-day mollusks and Yochelson, 

for the geologic data. Both contributed freely to the 

systematic work and conclusions.

ACKNOWLEDGMENTS

Type specimens of previously described species 

were generously lent for study by Roger L. Batten, 

American Museum of Natural History (AMNH), New 

York; Stig BergstrOm, Orton Museum, Department of 

Geology, Ohio State University (OSU), Columbus; 

and D. Alison, Redpath Museum, McGill University, 

Montreal, Canada. Additional materials were borrowed from the Redpath Museum and from the Illinois State Geological Survey, courtesy of Lois Kent.

We gratefully acknowledge the help and advice over 

the years of the following colleagues in assembling 

scattered literature and contributing geological and 

biological information: Michael Calver, Geological 

Survey of Great Britain, Leeds; Arthur Clarke, U.S. 

National Museum of Natural History (USNM), 

Washington, D.C.; Murray Copeland, Geological Survey of Canada, Ottawa; A. Cross, Western Michigan 

University, Kalamazoo; R. M. C. Eagar, University of 

Manchester, England; Laing Fergeson, Mount Allison 

University, Sackville, New Brunswick, Canada; S. M. 

Gasioroski and A. Gromczakiewicz, Polish Academy 

of Sciences, Krakow; John H. Hanley, U.S. Geological 

Survey (USGS), Denver; Radvan Horny, National 

Museum, Prague, Czechoslovakia; Russell Jacobson, 

Illinois State Geological Survey, Urbana; Hermann 

Jaeger, Humboldt Museum, Berlin; the late Boris Licharev, Ail-Union Geological Survey, Leningrad, 

U.S.S.R.; A. Logan, University of New Brunswick, 

Fredericton; J. P. E. Morrison, USNM; Bruce Runnegar, University of New England, Armidale, New South 

Wales, Australia; Thomas Phillips, University of Illinois, Urbana; Werner Vacecek, Krahuletz Museum, 

Eggenberg, Austria; the late Jon Weber, Pennsylvania 

State University; and Halina Zakova, Geological Survey of Poland, Kielce. Henry Berryhill and S. 

Schweinfurth, U.S. Geological Survey, guided the 

search for fossils in the Dunkard basin.

The SEM photographs were taken by Solem on a 

Cambridge S4-10 Stereoscan,2 purchased by the Field 

Museum of Natural History (FMNH), Chicago, with 

the aid of Grant 72-02149 from the National Science

2 Any trade names in this publication are used for descriptive purposes only and do 

not constitute endorsement by the U.S. Geological Survey.
STRATIGRAPHIC SETTING AND LOCAL OCCURRENCE

Foundation, whose assistance is gratefully acknowledged. The excellent photographic prints by Fred 

Huysmans were mounted for publication by Dorothy 

Karall and Elizabeth A. Liebman. Line drawings are 

by Elizabeth A. Liebman, Illustrator, Division of Invertebrates, Field Museum of Natural History. The 

assistance of Sharon Bacoyanis in library work and 

manuscript preparation and of Barbara Walden in 

compiling data was indispensible.

EARLIER SYSTEMATIC WORK

Dawson (1880) summarized previous work and 

added many new data. He treated six species from the 

following three localities: Coal-Measures at South 

Joggins, Nova Scotia (Pupa uetusta Dawson, 1867, 

Pupa bigsbii Dawson, 1880, and Zonites priscus Carpenter, 1867); Brian plant beds of St. John, New 

Brunswick (Strophites grandaeuus Dawson, 1880); 

and below coal No. 6 near Petty's Ford, Vermilion 

River, 111. (Pupa uermilionensis Bradley, 1872, Dawsonella meeki (Bradley, 1872). Whitfield (1881) 

named the genus Anthracopupa and described Anthracopupa ohioensis from near Marietta, Ohio; he also 

commented that Dawsonella had the aspect of the 

modern prosobranch family Helicinidae. Subsequently, Matthew (1895) described Pupa primaeva 

from the presumed Devonian Fern Ledges, Lancaster, 

St. John County, New Brunswick, and Stauffer and 

Schroyer (1920) described Anthracopupa(?) dunkardana, Naticopsis(?) diminuta, and Loxonema(?) 

parua from Permian rocks at Pleasant Grove, Belmont 

County, Ohio.

Starting with that of White (1883), several checklist summaries of Paleozoic land snails have been issued, and some systematic comments have been scattered in the literature, but they were not based on new 

information or new illustrations. Pilsbry (1926, p. 

316-319) reviewed the pupilloid forms. He recognized 

that Anthracopupa might belong to the Tornatellinidae or the Ellobiidae and described the genus Maturipupa for Pupa vermilionensis, suggesting that it too 

might be a tornatellinid. He also indicated that Dendropupa uetusta might be an ancestor of the Urocoptidae and stated that Pupa bigsbii, Pupa primaeva, 

and Strophites grandaeua could not be classified on 

the basis of pubished descriptions and illustrations. 

Henderson (1935) placed Strophites, Dendropupa, 

Anthracopupa, and Maturipupa in the Pupillidae, 

put Zonites priscus in the Zonitidae, and put Dawsonella in the Helicinidae. Yen (1949) essentially followed 

Henderson, except in considering that Zonites priscus 

and Strophites grandaeua were of uncertain affinity.

Bradley (1870, p. 254) reported the occurrence of 

two kinds of small land snails in "The nodular limemeeki, placing it next to the Helicinidae, had placed 

both Dendropupa and Strophella (=Strophites Dawson, 1880 not Deshayes, 1832) in a new subfamily of 

the Cyclophoridae, the Dendropupinae, and had 

placed Anthracopupa in a new subfamily, Anthracopupinae, of the Ellobiidae. Knight, Batten, and 

Yochelson (1960, p. 1279,1318) accepted the Dawsonellidae as a family unit, but referred the various pupilloid forms, including Anthracopupa and 

Maturipupa, to the Cyclophoridae, retaining the subfamily Dendropupinae Wenz (1938). H. B. Baker 

(1963) commented on the improbability of Anthracopupa and Maturipupa belonging to the Cyclophoridae. Zilch (1959-1960, p. 64-65) followed Wenz and 

placed both genera in a subfamily Anthracopupinae of 

the Ellobiidae.

STRATIGRAPHIC SETTING AND LOCAL 

OCCURRENCE

The bulk of our material comes from three widely 

separated areas. Each contains coal beds of economic 

importance, and interest in the coals probably led to 

the discovery of Paleozoic land snails. Each region is 

in a somewhat different geologic setting. The first, 

Vermilion County, 111., is on the west edge of the Illinois coal basin, where a repeated alternation of marine and nonmarine units first led to the concept of 

the cyclothem. The second region, the Dunkard basin 

is centered roughly on Belmont County, Ohio, and is 

characterized by lacustrine deposits. The third is the 

classic Joggins area of Nova Scotia and adjacent areas 

of New Brunswick, where an enormous thickness of 

terrestrial strata was deposited in a comparatively 

short time; interlayers of lacustrine sediments are present. The eastern Canada fossils are from strata dated 

as Westphalian B, equivalent to Middle Pennsylvanian age; the Illinois basin fossils are from the 

Kewanee Group of late Middle Pennsylvanian age; 

the Dunkard basin rocks are of Late Pennsylvanian to 

Early Permian age. By far the most fossil land snails 

are from the Dunkard basin, but this may reflect a 

lack of concentrated collecting for fossil land snails in 

the Illinois and eastern Canada areas.

Table 1 summarizes the general correlations between North America and Europe and indicates the 

approximate position of the stratigraphic groups that 

have yielded the specimens studied.

ILLINOIS AND INDIANA

Yen overlooked the fact that Wenz (1938, p. 435, 470) 

had erected a family Dawsonellidae for Dawsonella 

stone accompanying the fire clay of this seam * * *" at 

Petty's Ford on the Little Vermilion River, about four
NORTH AMERICAN PALEOZOIC LAND SNAILS, SUMMARY OF OTHER PALEOZOIC NONMARINE SNAILS

TABLE 1. General correlation of parts of the Pennsylvanian and 

Permian between North America and Europe, showing the approximate positions of the stratigraphic groups from which 

land snails have been collected in North America

Europe

PERMIAN

CARBONIFEROUS UPPER 

Autunian

Stephanian

Westphalian D

-i « 

^^(J *^

Westphalian B

Nova Scotia

Cumberland 

Group

Ohio

West Virginia 

Pennsylvania

Dunkard 

Group

Monongahela 

Group

Illinois 

Indiana

McLeansboro 

Group 1

Kewanee 

Group 1

North 

America

LOWER

1 _g

'?p S £<«

Missourian 

Series

Moinesian Des

Series

c 

& - 2 » < M

PERMIAN

PENNSYLVANIAN

As currently used in Illinois and Indiana nomenclature.

miles below Georgetown, 111. In his section, this fire 

clay with concretionary limestone was 5-20 feet 

(1.5-6 m) thick. Bradley (1872) indicated later that 

this was "* * * the concretionary limestone accompanying the underclay of coal No. 6 * * *," a well-known 

unit in the Illinois section. No. 6 coal is the "Herrin 

Coal Member"3 of the Carbondale Formation (Willman and others, 1975, p. 191). The Spoon Formation 

and overlying Carbondale Formation constitute the 

Kewanee Group. This group forms the lower part of 

the Des Moinesian Series in Illinois; the McLeansboro 

Group forms the upper part of the Des Moinesian and 

extends up into the Missourian.

Bradley's collection almost certainly came from 

above the "Vermilionville Sandstone Member"3 and 

was either from the "Big Creek Shale Member"3 or 

more likely from the upper part of the "Spring Lake 

Coal Member"3 of the Carbondale Formation, if those 

units extend eastward into Indiana.

1 As currently used in Illinois nomenclature.

In 1927, J. M. Weller obtained land snails along the 

Little Vermilion River, a short distance above Petty's 

Ford in SEV4 NWV4 sec. 2, T. 17 N., R. 11 W., Vermilion County, III; presumably these are topotypes of 

Bradley's two species. At that time the area was an 

abandoned strip mine. In 1976, no stratigraphic section could be compiled at this locality.

In his original description of Anthracopupa ohioensis, Whitfield (1881, p. 127) noted receipt of specimens of the two species described by Bradley (1872) 

from John Collette, then State Geologist of Indiana. 

He later amplified this remark slightly (Whitfield, 

1895, p. 490), stating "* * *specimens of the two forms 

from that State * * *," implying that the land snails 

came from Indiana. In a review of Whitfield types 

purchased by the University of California, Peck and 

McFarland (1954, p. 304, 306) recorded Dawsonella 

meeki (Bradley) and Pupa vermilionensis Bradley 

from "Carboniferous, Coal Measures, Little Vermillion River, Vermillion Co., Indiana"; the county name 

in Indiana has a double "1," but the river and Illinois 

county name have one "1."

In view of Whitfield's remark in 1881, the specimens now in California are probably those he referred 

to as being from Indiana, and probably they do not 

have the status of types. One specimen of each of 

Bradley's species was illustrated by Peck and McFarland (1954). If the specimen of Dawsonella meeki that 

they photographed is the same as that figured by 

Whitfield (1881, figs. 5, 6), then Whitfield did not accurately portray its features.

To the best of our knowledge, Whitfield's remark is 

the only published report of Paleozoic land snails 

from Indiana. Confirmation that Paleozoic land snails 

are in Indiana was substantiated by J. M. Weller who 

made a collection from SWV4 , sec. 29, T. 17 N., R. 10 

W., Vermillion County, Ind., years ago. In attempting 

to recover this locality, Russell Jacobson found another occurrence of gastropods in a strip coal mine 

along the Little Vermilion River in the center NEV4, 

sec. 32, T. 17 N., R. 10 W., Vermillion County, Ind. 

The limestone is about 15 cm thick, nodular, and brecciated. Gastropods do not appear on the surface but 

are seen abundantly when the material is sawed. The 

limestone is about 1.5 m below a coal thought to be 

coal no. 4 (Summum coal), and the limestone is designated the equivalent of the Breezy Hill Limestone 

Member of the Cabaniss Formation of Kansas (Shaver and others, 1970, p. 229). If the coal is correctly 

identified, this new locality is in the Petersburg Formation of Indiana usage and is slightly older than the 

occurrence in eastern Illinois.

Collections from Indiana and Illinois contain several times as many fossils of Dawsonella meeki (Brad-
STRATIGRAPHIC SETTING AND LOCAL OCCURRENCE

ley) as of P. vermilionensis. The limestone matrix 

appears slightly more fine grained than in most of the 

gastropod-bearing beds of the Monongahela and Dunkard Groups, though this may be a function of the degree of weathering.

Terrestrial gastropods, known from two other localities in Illinois, have never been documented. These 

fossils suggest that a study of the freshwater limestones of the Illinois basin would significantly increase 

the number of localities known to contain fossil terrestrial gastropods; such freshwater limestones tend to 

be neglected.

The first locality is in the center NVfc sec. 27, T. 7 N., 

R. 10 E., Jasper County, 111. The fossils are from an 

unnamed freshwater limestone about a meter below 

the "Reisner Limestone Member of the Mattoon Formation."4 That marine limestone is near the base of 

the "Mattoon Formation,"4 and the formation is the 

youngest in the McLeansboro Group. Two poorly preserved Anthracopupa specimens and abundant specimens of Spirorbis were collected from this locality 

long ago.

The second locality, also one from which fossils 

were collected long ago, is on the south bank of 

Hodges Creek, northwest of Chester Field, SW1^ sec. 

30, T. 10 N., R. 9 W., Macoupin County, 111. A field 

check indicated that this area is badly overgrown. The 

stratigraphic section is thinner and contains more marine units than the section in the eastern margin of the 

Illinois basin. Possibly the fossils come from a freshwater limestone in the greatly thinned interval between coal no. 6 (Herrin) and coal no. 4 (Summum) or, 

less likely, they come from below the Summum coal. 

These fossils are thus middle Des Moinesian in age. 

Poorly preserved Anthracopupa occur at this locality.

OHIO, PENNSYLVANIA, AND WEST VIRGINIA

The Dunkard basin is west of the Allegheny Mountains and is more or less bisected by the Ohio River. 

Erosion has truncated the margins; the basin may 

have extended to western Maryland as well as an unknown distance northeastward. It is a broad syncline 

in which rocks are essentially flat lying. Above the marine Conemaugh Group, of Late Pennsylvanian age,, 

are the Monongahela Group, also considered Late 

Pennsylvanian in age, and the Dunkard Group, considered by various workers to be Pennsylvanian or 

Permian (Barlow, 1975). The strata in the upper half 

of the Dunkard, the Greene Formation, are considered 

by most workers to be Early Permian.

Exposures are moderately good in the Dunkard basin, and no major structural complications exist there.

4 As currently used in Illinois nomenclature.

Because marine strata are absent in the Monongahela 

and younger beds, a great deal of reliance is placed on 

the various coal beds for correlation. Although a few 

miscorrelations in the past resulted from misidentification of coals in widely spaced sections, detailed 

mapping and more closely spaced sections (Cross and 

others, 1950; Cross and Schemel, 1956; Berryhill, 

1963; Berryhill and others, 1971) have essentially 

eliminated stratigraphic errors of this sort.

About 1,000 m of strata is exposed in the Dunkard 

basin; it consists of coal, sandstone, freshwater limestone, and shale of several varieties. The area is readily interpreted as a swamp-lake complex into which 

rivers debouched. From southwest to northeast across 

the basin, deposits generally shift from terrestrial to 

aquatic. Because it has the thickest and most abundant limestone deposits, Belmont County, Ohio, appears to have been the most aquatic part of the basin.

The unit that has yielded the most gastropods is the 

Little Captina Limestone Member of the Monongahela Formation. This unit has been examined at various localities throughout 3,885 km2 of outcrop, and at 

most places it yields gastropods. The characteristic sequence of this member is a thin rubbly limestone a few 

centimeters thick at the bottom, then about 15-20 cm 

of shale, a massive more fine grained limestone nearly 

a meter thick, a shale, and a thin rubbly limestone on 

top. The thin limestones above and below the main 

bed have yielded specimens.

The lower limestone member of the Washington 

Formation in the lower half of the Dunkard Group is 

the source of the second largest number of fossils. In 

this member of the Washington, as in the Little Captina Limestone Member of the Monongahela, the rubbly limestone, not the main limestone unit, ha» 

yielded fossils. Perhaps the main limestone unit only 

appears unfossiliferous because it has not been as 

weathered as the rubbly limestone. Some of the rubbly limestones might be brecciated limestones formed 

from drying crusts of lime mud.

Yochelson (1975, p. 250) suggested that A. ohioensis 

was probably from the Little Captina Limestone 

Member of the Monongahela, although its origin in 

the Washington Formation cannot be ruled out. In either event, collections in the Field Museum of Natural 

History provide material from the general type area at 

both horizons.

The three species of gastropods, Anthracopupa (?) 

dunkardana, Loxonema (?) parva, and Naticopsis (?) 

diminuta, from the Dunkard Group that were described by Stauffer and Schroyer (1920, p. 144-145) 

all were obtained "from the shales at the base of the 

Lower Washington Limestone, one-half mile south of 

Pleasant Grove, Belmont County, Ohio." Yochelson
NORTH AMERICAN PALEOZOIC LAND SNAILS, SUMMARY OF OTHER PALEOZOIC NONMARINE SNAILS

visited this area several times without finding an outcrop of the "Lower Washington Limestone Member." 

Material from U.S. Geological Survey collections 

24846-PC and 24855-PC is from the top of this unit, 

about a kilometer from the type locality.

NEW BRUNSWICK AND NOVA SCOTIA

More confusion has surrounded the age of land 

snails from Joggins, Nova Scotia, and St. John, New 

Brunswick, than has surrounded the age of those from 

the Illinois and Dunkard basins. The gastropods at 

Joggins were first found by J. W. Dawson and Charles 

Lyell in 1851. For decades they were considered to be 

as old as Devonian. The same illustration and figures 

were repeated by Dawson (1855,1868) in various editions of "Acadian Geology" and elsewhere during the 

last century, but, since the specimens were first described, no further study of the specimens has been 

made. The one exception to this is the redescription 

and photographic reillustration by Knight (1941, p. 

98-99, pi. 96, figs. 6a-6e) of Pupa vetusta, the type 

species of Dendropupa.

Dendropupa vetusta ranges locally through more 

than 610 m (2,000 ft) of the Cumberland Group. 

The lowest bed in which Pupa vetusta occurs belongs to group VIII 

of Division 4 of my section of the South Joggins, and is between 

Coal 37 and Coal 38 of Logan's section, being about 42 feet below 

Coal 37. The next horizon, and that in which the shell was first 

discovered, in 1217 feet of vertical thickness higher, in group XV of 

Division 4 of my section. The shells occur here in erect Sigillariae, 

standing on Coal 15 of Logan's section. The third horizon is in 

group XXVI of Division 4, about 800 feet higher than the last. Here 

also the shells occurred in an erect Sigillaria (Dawson, 1880, p. 

408-409.)

The collection at the Redpath Museum contains 

several hundred fragments of Dendropupa, all in 

dark-gray carbonaceous shale. The original occurrences of D. vetusta in hollow upright tree stumps 

suggest that these were preserved in place, quite unlike the floated shells in lacustrine limestones. Sedimentation was intermittent but rapid when it took 

place. Upright trees 9 m (30 ft) high are preserved 

(Poole and others, 1970, p. 294). Much of the Cumberland Group, especially the upper half, is relatively 

coarse.

Although several paleontologists have studied the 

fossils of the Cumberland Group, emphasis has been 

placed on the fossil plants. Bell (1944, p. 29) did note 

in passing that Dendropupa occurs in the Cumberland Group, of late Westphalian A or early Westphalian B Age. Copeland (1957, p. 14) assigned the 

Cumberland to Westphalian B.

Hacquebard (1972) correlated the Cumberland to 

Westphalian B with a high degree of confidence principally on the basis of spores. He also summarized the 

depositional history of the area and included an interpretive sketch showing the coal basins of northern Europe and eastern Canada as adjacent. Poole and 

others (1970, Chart II) correlated the Cumberland as 

late Westphalian A and Westphalian B, in part Atokan in the American sense.

In addition to Dendropupa vetusta, Dawson (1880) 

reported three land snails from the Joggins section. 

The form he referred to as "variety tenuistriata" occurred "Along with the ordinary form" of D. vetusta. 

He does not state clearly whether he differentiated 

this variety at all three stratigraphic levels where the 

species was collected. Except for a drawing of part of 

the shell surface, he did not illustrate the variety.

The second form, Pupa bigsbii, was named by Dawson (1880, p. 410) for a few specimens "*** found in 

the erect trees of group XV at the Joggins, along with 

D. vetusta" Only one specimen was available for 

study, and it was free of matrix. We accept Dawson's 

statement of occurrence; presumably this form also 

was found essentially where it had lived, in a decaying 

tree which was covered by an influx of sediments.

The final species from Joggins, Zonites (Conulus) 

priscus Carpenter (1867), was recorded by Dawson 

(1880, p. 411, 412) as occurring in the lowest beds at 

South Joggins that produced D. vestusta (Group 

VIII). Upon excavation of the locality in an attempt to 

find a greater variety of fossils, Dawson found "* * * 

that the thin layer containing the land shells was not 

continuous but in limited patches." He found only a 

few Zonites and a bone fragment in the layer. "One or 

two crushed specimens have been subsequently found 

in the erect trees holding Pupa vestusta in group XV, 

but the species is extremely rare in comparison" 

(Dawson, 1880, p. 410). On the basis of the meager 

data supplied, we suggest that this might have been a 

thin lacustrine limestone of a local pond into which 

the snail shells floated from their habitat in and 

around large trees.

Land snails are exceedingly rare in the Fern Ledges 

at St. John, New Brunswick. No systematic reports of 

Paleozoic land snails in eastern Canada have been 

published since the report published by Matthew in 

1895. In part, this lack of reports is probably due to 

geologists ignoring the snails because they are of little 

stratigraphic value. The land snails received a great 

deal of publicity during Dawson's lifetime; the publicity may have served to defer further investigation of 

them.

The age of gastropods in the Little River Group 

from "Plant bed No. 2" of the Fern Ledges at St. John, 

New Brunswick, is more poorly understood than is the 

age of those from the Joggins Section. The gastropods 

in the Little River Group were originally considered 

Devonian (Erian) and subsequently were reassigned
ANALYSIS AND CLASSIFICATION OF TAXA

to the Silurian. Stopes (1914, p. 3-8) has summarized 

the various age assignments for these beds. The Fern 

Ledges are now placed in the Little River Group of 

late Carboniferous age. Correlation of the stratigraphic units in the nonmarine basins of eastern Canada 

is extremely difficult, but apparently the flora is comparable with that of the lower part of the Kanawha 

Formation in West Virginia. If this correlation is correct, the beds at Fern Ledges are probably of Westphalian B Age and therefore virtually the same age as 

those of the Joggins area that yielded gastropods. The 

section at Fern Ledges is thinner than that at Joggins.

Pupa primaeva Matthew (1895, p. 99) was collected 

from "Dark gray shale of Plant Bed No. 2. Fern 

Ledges, Lancaster, St. John County, N.B." Repeated 

unsuccessful attempts have been made to trace Matthew's material. One can infer from the description 

that only a single specimen was collected.

Matthew's (1895, p. 99) species "comes from the 

plant bed in which many years ago, the author found 

the species of Pulmonate described as Strophites 

grandaeva by J. W. Dawson." Strophella grandaeva 

(Dawson) is known from a single flattened specimen 

and is discussed on p. 14.

ANALYSIS AND CLASSIFICATION OF TAXA

Zonites (Conulus) priscus Carpenter, 1S67

Plate 2, figures 1-5; plate 8, figures 1-2

Zonites (Conulus) priscus Carpenter, in Dawson, 1867, p. 331-333, 

figs, a-c, e; Dawson, 1868, p. 385, fig. 150; Dawson, 1880, p. 411- 

413, figs. 10,11,14d; Yen, 1949, p. 237; Yochelson and Saunders, 

1967, p. 246.

Zonites priscus Carpenter, White, 1883, p. 453, 475, pi. 2, figs. 11- 

12; Henderson, 1935, p. 155.

Pyramidula priscus (Carpenter), Pilsbry, 1895, p. xxxix.

Archaeozonites priscus (Carpenter), Grabau and Shimer, 1909, p. 

822, figs. 120h-i.

No meaningful discussion of the systematic position 

of Zonites priscus has been given, except the brief 

comment by Pilsbry (1895, p. xxxix) that this species 

should be compared with Discus (= Pyramidula of 

1895 usage) or Charopa. The suggestion by Yen (1949, 

p. 237) that this species might be a cyclophorid is unfortunate and relates to Yen's misinterpretation of 

Dawson's original drawings. Dawson's illustrations 

are excellent but included several assumptions as to 

structure that are not supported by SEM observations.

We were able to examine six specimens, three of 

which are illustrated here on plates 2 and 8. The specimen illustrated by Dawson in 1867 (fig. b) and in 1880 

(fig. 10) as a whole-shell view was designated as the 

holotype by Alison and Carroll (1972); it is catalogued

as Redpath Museum 3116 (pi. 2, fig. 1). This specimen 

has 3V4 whorls. Despite being crushed laterally, it is 

2.63 mm in diameter. A paratype, Redpath Museum 

3116a (pi. 8, figs. 1,2), is the specimen used by Dawson 

(1867, p. 332, fig. e; 1880, p. 411, fig. 11) for the basal 

view. The maximum diameter of this specimen is 3.22 

mm. Dawson's figures of the shell base do not agree, as 

the earlier drawing (Dawson, 1867, p. 332, fig. e) shows 

that specimen with an extended columellar area and 

open umbilicus, whereas the later (Dawson, 1880, p. 

411, fig. 11) shows the columellar lip joining the preceding whorl without any trace of an umbilical opening. Examination of the specimen shows that the 

basal, columellar, and parietal areas have been destroyed and that the fossil is an internal mold (steinkern). Hence, no information is available concerning 

the exact structures in these areas. The third specimen (Redpath Museum 3116b, pi. 2, figs. 2-5) is a juvenile paratype having 2V4 whorls; it is only 1.94 mm 

in diameter. The excellent preservation of this fossil 

permitted illustration of shell-structure details (pi. 2, 

figs. 2-5) and determination of the species affinities.

The other three examples are not illustrated. The 

specimen labeled "upper bed" is 2.43 mm in diameter 

but is badly crushed, the specimen labeled "erect tree" 

is too crushed to measure, and the final specimen is an 

external impression of a short section of the body 

whorl which shows details of sculpture but no other 

features. Presumably all three specimens represent 

paratype material.

The SEM photographs indicate that early interpretations of the sculpture were erroneous. Both P. P. 

Carpenter (in Dawson, 1867, p. 332, fig. c) and Dawson 

(1880, p. 411-412, fig. 14d) reported that the apical 

sculpture was continued onto the postapical whorls 

and that the major ridges had four to five microradial 

ridges between each pair. Because of the small size of 

the shells, the limits of optical equipment and illumination, and the lack of knowledge concerning the complexity of shell microsculpture that existed at that 

time, these early conclusions were reasonable approximations.

The apical sculpture (pi. 2, figs. 2-4) is distinctly 

different from the postapical. At very low magnification (pi. 2, fig. 1), the apical sculpture does appear irregularly radial, and when it is viewed from a low 

angle (upper right of pi. 2, fig. 2 and lower left of pi. 2, 

fig. 3), its obliquely radial aspects are emphasized. 

When viewed from an angle closer to the vertical (center and lower part of pi. 2, fig. 2 and pi. 2, fig. 4), the 

radial elements show an irregular rugose aspect. The 

main radial elements are partly interrupted at seemingly regular intervals. The difference between this 

sculpture and the postapical sculpture is quite clear.
8 NORTH AMERICAN PALEOZOIC LAND SNAILS, SUMMARY OF OTHER PALEOZOIC NONMARINE SNAILS

The postapical sculpture consists of major radial ribs; 

most commonly, two or three microradial ridges are 

between each pair of major ribs (pi. 2, fig. 2 right and 

especially pi. 2, fig. 3 lower center). In addition, when 

viewed at high magnification and from an oblique angle (pi. 2, fig. 5), the sculpture in the suture suggests 

that a microspiral element probably was present. Particularly apparent in the third and fourth interstices 

from the bottom of the photograph are remnants of 

spiral ridges at regular intervals. The sutural sculpture of heavily sculptured shells characteristically 

shows least wear, even when other parts of the shells 

are significantly worn. Preservation of sculpture in 

this section of the body whorl suture is a fortunate accident, but it is not unusual. Recent shells that show 

an equivalent amount of wear have sculptural remnants of no greater prominence.

On the basis of these observations, the following 

points are critical in attempting to determine affinities: the apical and postapical sculptures are quite 

distinct from each other; the apical sculpture has a 

major oblique radial component that is interrupted 

over most of the apex; the postapical sculpture consists of major radial ribs and a small number of microriblets in between; and apparently a spiral 

microsculptural element is present on the postapical 

whorls, at least near the suture.

Although no comprehensive review of sculpture 

patterns in extant land snails exists, the combination 

of different apical and postapical structures and the 

presence of major sculpture and microsculpture on 

the postapical whorls fits the "endodontoid" complex 

of families. Zonitoids generally lack apical sculpture, 

except fine spiral lines or ridges, rarely have prominent major radial sculpture, and essentially never 

show microradial sculpture. Pupilloid taxa generally 

have apices that are smooth or have microspiral 

grooves, followed by postapical sculpture at most major radial ribs, even in the planulate taxa. The bulimulids, which are much larger and generally elongated, 

do have complex apical sculpture and a variety of 

post-apical sculptures, but apparently they do not 

have a combination of macroradial and microradial 

postapical sculpture.

Several minute helicoid taxa that at one time were 

classified as endodontoids have been shown by dissection to belong to other families, despite having combined macroradial and microradial postapical 

sculpture and distinctive apical sculpture. Solem 

(1977) reviewed the sculpture of Striatura, Punctum, 

Radiodiscus, and Planogyra, demonstrating differences in formation and detail that separate them from 

the endodontoid families Charopidae and Endodontidae, which have the most complex and conservative

sculpture patterns of the endodontoid complex. The 

Endodontidae have a combination of major radials, 

microradials, and "squiggly" spiral sculpture on the 

apex (see Solem, 1973b, fig. 1; 1976) and have macroradial ridges, microradial ridges and minute spiral 

ridges on the postapical whorls. The Charopidae 

primitively have an apical sculpture of spiral cords, 

which subsequently may be overlaid by added radial 

elements; they also have a characteristic, very complex postapical sculpture of major radials, microradials, and spiral elements that buttress the 

microradials (see Solem, 1970, figs. 7-11). The Punctidae show a variety of elements, but normally these are 

periostracal in nature, are semideciduous, and rarely 

show microspiral elements; the apical sculpture most 

frequently consists of spiral elements only.

Previously, little attention has been paid to the 

sculpture in the Discidae, a Holarctic family. The 

sculpture in this family provides the key to understanding the apical sculpture and affinities of Zonites 

priscus. The limits of the family Discidae are defined 

elsewhere (Solem, 1978a). Pilsbry "(1948, p. 566, fig. 

303) pointed out that some discids had complex apical 

sculpture, but his illustration did not show their complexity. The sculpture of Anguispira picta (Clapp, 

1920) is illustrated in plate 3; this species has marked 

apical sculpture, but the postnuclear sculpture is lost 

on the penultimate and body whorls (see Pilsbry, 

1948, p. 589). At low magnification (pi. 3, fig. 3), the 

basic pattern of change from dominance of microelements on the early apex to the presence of only major 

radial elements on the postnuclear whorls is evident.

Examined at higher magnification, the details of 

this change can be worked out. The upper part of the 

apex (left third of pi. 3, fig. 1) has diagonal radial microriblets that at a later stage in life (lower left of pi. 3, 

fig. 5) begin to be crossed by oblique incised lines, creating the checkerboard pustule effect seen on plate 3, 

figure 5. Near the tip of the apex (pi. 3, fig. 4, far left), 

the initial irregular anastomosing nature of the microradial sculpture can be detected. Shortly before 

the end of the first apical whorl (pi. 3, figs. 3, 5), an 

overlay of prominent radial ribs begins, starting initially at the upper suture and extending not even to 

the midpoint of the exposed whorl surface. After the 

first dozen or so of these ridges, they reach all the way 

to the next suture. At the 1 Vi -whorl mark (pi. 3, fig. 4), 

these radial ridges are sharply defined and very 

prominent. The microsculpture between the radial 

ridges has been slightly altered (pi. 3, fig. 6), the oblique element being greatly reduced in prominence. 

By the 1 Vz -whorl point, a second shift has taken place; 

the oblique microsculpture once again is very prominent, and the pustules are quite marked (pi. 3, fig. 2).
ANALYSIS AND CLASSIFICATION OF TAXA

The apical sculpture in this species thus is composed of three basic elements, whose prominence 

shifts from one part of the apex to another. Initially, 

anastomosing, strongly retractive microradial riblets 

having very narrow interstices are present. These become clearly separate entities but are then crossed at 

essentially right angles by extremely protractive incised lines, turning the radial riblets into a checkerboard of pustules. Superimposed on this pattern is a 

rather gradual development of protractive radial ribs 

that are inclined at about a 45° angle and are slightly 

curved. When this third element reaches full size, the 

oblique microelements are reduced in prominence, 

only to reappear near the end of the apex in full 

prominence.

The early postnuclear sculpture (pi. 3, fig. 1 right, 3, 

4 far right, 5 left) consists of low rounded radial ribs 

that essentially are continuations of the superimposed 

apical radial elements, plus a few microradial riblets 

in between the major ribs.

Except for the third element of the apical sculpture, 

the radial ridges, and the absence of any microspiral 

postapical sculpture, the shell sculpture of Anguispira picta agrees with that of Zonites priscus. The 

pattern of interruptions to the oblique radials in the 

latter species (pi. 2, fig. 4) can be interpreted as the 

remnants of dissecting spiral lines under conditions of 

moderate wear. Thus, the sculptural elements of Zonites priscus are identical with the basic pattern found 

in extant Discidae.

Because of the limited material and the fact that 

several specimens were crushed, the adult size and 

shape of Zonites priscus are difficult to estimate. The 

whorls of the holotype (pi. 2, fig. 1) increase rapidly in 

width. Their pattern resembles the whorl incremental 

pattern found in the modern New Zealand Flammulina von Martens, 1873, rather than the more regular, 

less rapid incremental patterns of the North American genera Discus Fitzinger, 1833, and Anguispira 

Morse, 1864. The sculpture on the last section of the 

body whorl in the holotype (pi. 2, fig. 1, right) is both 

crowded and irregular, which is typical of gerontic 

growth in most land-snail taxa. The major radials are 

close together and are somewhat irregular in shape, 

and no microradials are clearly visible. This specimen 

has 3V4 whorls; thus, both the whorl count and the 

whorl incremental pattern of Zonites priscus are comparable with those of Flammulina. Because of great 

lateral crushing, the measured diameter of the holotype is only 2.63 mm, whereas the measured diameter 

of the steinkern (pi. 8, figs. 1, 2) is 3.22 mm. Probably 

Zonites priscus had a maximum shell diameter of 

about 3.5 mm and as many as 3 Vi whorls.

On the basis of the above data, we have no hesitation in assigning Zonites priscus to the Discidae and 

placing it in a new genus, described below.

Genus Protodiscus Solem and Yochelson, new genus

Type species.—Zonites (Conulus) priscus Carpenter, 1867.

Diagnosis.—Small, diameter 3-3.5 mm, shells having a relatively loose coiling pattern, whorl count 3 1A 

to 3V2, apex distinctly elevated. Apical sculpture of 

protractive radial ribs, probably interrupted by equally strongly retractive lines, appearing rugose when 

worn. Postapical sculpture of low, prominent, protractive radial ribs, two or three microradial riblets between each pair of major ribs, and microspiral 

ridgelets visible near the suture.

Discussion.—The name refers to the shell ornamentation that serves as a prototype for the sculpture 

found in modern Discidae. The fact that the apical 

ridges in Protodiscus are protractive, whereas those in 

the modern Anguispira are retractive, is not significant, in view of the shifting sculpture seen on the apex 

of Anguispira (pi. 3). The age and occurrence of the 

type and only known species were discussed on p. 6.

Pupa bigsbii Dawson, 188O

Plate 2, figure 6

Pupa bigsbii Dawson, 1880, p. 410, figs. 5, 6; Pilsbry, 1926, p. 318.

Pupa bigsbyi Dawson, White, 1883, p. 453, 456, 475, pi. 2, 

figs. 9, 10.

Dendropupa? bigsbii (Dawson), Henderson, 1935, p. 149.

Anthracopupa? bigsbii (Dawson), Yen, 1949, p. 235-238; Yochelson and Saunders, 1967, p. 31.

The two specimens on which this taxon were based 

have not been studied since their original description. 

We select as lectotype the larger specimen that was 

illustrated in figure 5 in Dawson (1880). It is catalogued as Redpath Museum 3122. The lectotype is 

2.19 mm high and 1.18 mm in diameter and has about 

4V2 whorls remaining. The specimen is covered in 

back with a hard glue, but the front side is unobscured 

and is illustrated on plate 2, figure 6. No trace of surface sculpture exists. Both the shell lip and the last 

part of the body whorl are completely missing. The 

part of the specimen farthest from the apex is a piece 

of shell, presumably part of the columellar lip; the 

shell height has been measured to this point. The 

specimen is very slightly flattened but probably not 

enough to distort the form significantly.

The very broad apex, curved spire contour, and sudden change to the more flattened left profile of the 

body whorl, are all features that are characteristic of 

the modern pupillids (see discussion of Anthraco-
10 NORTH AMERICAN PALEOZOIC LAND SNAILS, SUMMARY OF OTHER PALEOZOIC NONMARINE SNAILS

pupa and figure 6A). Species such as Gastrocopta tappaniana (Adams, 1842), G. pentodon (Say, 1821), 

various Papilla, and particularly Vertigo (see Pilsbry, 

1948, p. 887, fig. 477; p. 930, fig. 502; p. 945, fig. 509; p. 

958, fig. 515) match the basic whorl contours. This 

pattern of spire shape is highly characteristic of the 

Pupillacea but is not found in the Tornatellinidae. 

Thus, we have no hesitation in placing Pupa bigsbii in 

the Pupillacea. Without knowing the apertural characters, we cannot assign it to any generic taxon. We 

consider that too little information is available to warrant establishing a new genus for it. We have no 

doubts that it does represent a distinct generic unit, 

but we cannot characterize a genus adequately.

Another specimen studied by Dawson (1880, fig. 6) 

is catalogued as Redpath Museum 3122a. It is imbedded in matrix and measures 1.97 mm in height and 

1.28 mm in diameter. Careful cleaning along one edge 

and a low-angle photograph (pi. 8, fig. 3) revealed 

traces of low radial ribs on the shell. On plate 8, figure 

4, the whorl contours can be seen more clearly. The 

narrower apex, less curved spire, and presence of radial sculpture are very different from the characteristics 

of the lectotype. This specimen is an Anthracopupa, 

not "Pupa" bigsbii. We chose not to name it as a distinct species because it is a juvenile shell in a poor 

state of preservation, and the apertural features are 

unknown. However, the specimen is significant, for no 

other Anthracopupa are known from the Joggins, and 

this unique individual extends the range of the genus 

downward.

One additional uncatalogued specimen in the Redpath Museum is badly crushed and broken in two, 

then crudely glued back together. It appears to be 

"Pupa" bigsbii. Because no apertural features are discernible, it adds nothing to the concept of the species.

Dendropupa vetusta (Dawson, 1855)

Plate 4; plate 8, figure 5; figures la, b

Pupa vetusta Dawson, 1855, p. 160, fig. 12; Dawson, 1860, p. 270,

figs. 1-3; Dawson, 1880, p. 405-409, figs. 1-4, 14, a, b; White,

1883, p. 453, 456, 475, pi. 2, figs. 1, 2.

Pupa vetusta variety tenuistriata Dawson, 1880, p. 406, fig. 14b. 

Dendropupa vetusta vetusta (Dawson), Henderson, 1935, p. 149,

150. 

Dendropupa vetusta tenuistriata (Dawson), Henderson, 1935, p.

150; Yochelson and Saunders, 1967, p. 65. 

Dendropupa vetusta (Dawson), Pilsbry, 1926, p. 317, 318, pi.

32, figs. 21-24; Yen, 1949, p. 235, 238; Knight, 1941, p. 98, pi.

96, figs. 6a-e; Yochelson and Saunders, 1967, p. 64.

This species has been variously called a pupillid or a 

urocoptid similar to Microceramus (Dawson, 1880, p. 

408; Pilsbry, 1926, p. 318), but no assured systematic 

position has been established. The data shown on 

plates 4 and 8 permit assignment to a family unit, although *with less assurance than for Anthracopupa 

and Protodiscus.

The holotype of Dendropupa vetusta was designated by Knight (1941, p. 98). Three figured specimens 

at the Redpath Museum that are catalogued under 

3119 are incorrectly designated as cotypes. The material we have examined has not been figured previously 

and was part of Dawson's uncatalogued collections. 

Though several hundred individuals are present, almost all are fragments. The material was excellent for 

studying sculpture and shell structure, but no quantitative study of size variation was possible.

The apex of the shell in this species (pi. 4, fig. 1) is 

about 2 Vz whorls in length, and the whorls are evenly 

rounded. It is not bulbous, and there is no significant 

change in logarithmic spiral. The sculpture on the apical whorls (pi. 4, fig. 2, lower right) consists of very 

fine radial incised lines that are quite crowded and follow the same angle as the growth wrinkles. Dawson 

(1880, p. 405-406) interpreted this sculpture as follows: "Nucleus round, smooth, the first turn below the 

nucleus marked with rows of little pits which gradually pass into the continuous striae." The worn areas 

having pitted spots on the surface of many specimens 

could easily be interpreted in this way. This observation by Dawson has been cited by most people who 

commented on the species and has greatly influenced 

subsequent speculations as to its affinities. We inspected several specimens that are quite well preserved but failed to detect any sculpture on the apex 

other than the very fine and crowded radial lines.

The postapical sculpture consists of relatively 

prominent radial ribs (pi. 4, fig. 2) that vary somewhat 

in size and spacing. Individual ribs, as seen in a fracture section (pi. 4, fig. 4), are shaped like broad inverted "V's" and extend from suture to suture. At 

their origin in the upper suture (pi. 4, figs. 2, 3), the 

ribs are blunt tipped and have no sign of expansion or 

crenulation effect. No microsculpture is visible, even 

at high magnifications. On the last part of the body 

whorl, the radial sculpture is reduced in size and quite 

crowded, almost disappearing behind the lip (pi. 4, fig. 

6). The lip itself is broadly and gradually flared, as was 

well shown in Dawson's figure 2 (1880, p. 407). Seen 

from behind (pi. 4, fig. 6), the gradual expansion of the 

lip is evident. A drawing of an apertural fragment (fig. 

IB) also illustrates the broad and gradual lip expansion. There is no heavy callus on the parietal wall or a 

continuous lip across the wall.

Preservation of detail is good in many specimens. 

The original shell material in the figured specimen has 

been replaced (cross section on pi. 4, fig. 4) by a single 

layer of crystals extending completely across the 

whorl profile. At places (pi. 4, fig. 5), highly unusual
ANALYSIS AND CLASSIFICATION OF TAXA 11

2mm

2mm

FIGURE 1. Shell structures in Dendropupa and a modern enid. AB, Dendropupa uetusta (Dawson, 1855). Coal Measures, South 

Joggins, Nova Scotia. A, Polished section through partly crushed 

specimen. Arrows indicate columellar ridges; a large shell axis 

cavity is shown. Redpath Museum 15.024. B, Lip from another

crystal patterns are on the surface of the shell-replacement material. In other specimens (pi. 4, fig. 6), original shell material seems to have been retained, and 

shell-layer structures have been preserved. No study 

of the shell material was attempted, because knowledge of shell layers in comparable Holocene taxa is too 

inadequate to make meaningful comparisons.

The data obtained from an old polished cross section of a shell in the Redpath (fig. 1A) and study of an 

isolated columellar fragment (pi. 8, fig. 5) proved crucial in restricting the options for classifying D. uetusta. In figure 1A, only actual shell fragments are 

shown. The specimen had been partly crushed, and 

the polishing was done at an angle. Only the lower 

part of the columella is present; the upper columellar 

part was either lost before preservation because of 

crushing or, more probably, was ground away. We 

have no data concerning the diameter of the columella 

in the upper whorls in D. vetusta. A steinkern of the 

European D. zarecznyi Panow, 1936 (see p. 27) has the 

open columella confined to the lower half of the spire. 

We suspect this may be the situation in D. vetusta. 

The shell was polished from the back side, and thus 

the aperture would have been shown on the far left of 

figure 1A.

2mm

fragment, showing broad expansion, the thinness of the expansion, and pattern of junction to penultimate whorl. Redpath Museum 15.032. C, Columellar ridge in a specimen of Buliminus 

labrosus labrosus Olivier, 1804. FMNH 54038.

The significant feature is that Dendropupa has a 

very wide, hollow columella, on which there are two 

raised ridges (arrows in fig. 1A). The ridge visible on 

the left side of the diagram is lower on the whorl profile than the one shown on the right side. The ridge on 

the right side also can be seen as a remnant on the 

next higher whorl on the left side. The SEM photograph (pi. 8, fig. 5) confirms that two such barriers existed and gives some idea as to their shape and angles 

in relation to the shell axis. Neither barrier apparently 

would have been visible from the aperture. Their presence had been postulated by Knight (1941, p. 98).

The internal barrier ridges on the columella are sufficient to eliminate possibility of placing Dendropupa 

in any prosobranch group such as the West Indian Po-^ 

matiasidae or the elongate cyclophoroid taxa. The 

presence of an operculum capable of shutting the shell 

aperture and the developed ridges on the interior of 

the columella are not compatible features. The small 

toothlike ridge or slit at the aperture in many cyclophorids and diplommatinids is a device to permit air 

flow past a closed operculum (see Solem, 1974, p. 178, 

179) and in some species is intensified to form an actual breathing tube. These are very different adaptations from having on the columella actual long ridges 

that would prevent free movement of an operculum.
12 NORTH AMERICAN PALEOZOIC LAND SNAILS, SUMMARY OF OTHER PALEOZOIC NONMARINE SNAILS

Only higher mesogastropods and neogastropods having reduced opercula have columellar ridges. For this 

reason alone, the suggestion of Wenz (1938, p. 470) 

that Dendropupa might be a cyclophorid has to be 

abandoned.

Comparison with extant pulmonate families is required. Several families characteristically have elongated shells that are roughly equivalent to 

Dendropupa in shape and size. The Clausiliidae and 

most of the Pupillacea differ in having complex apertural barriers as adults or near adults. These barriers 

are clearly visible from the aperture and frequently 

edge the aperture itself. The Tornatellinidae differ in 

being generally smaller than Dendropupa, and, except in the Hawaiian radiation, the species have complex apertural barriers at all stages of growth. The 

Subulinidae and Achatinidae, except very rare individuals, lack a reflected lip, have different patterns of 

shell growth than Dendropupa, practically never have 

continuous radial ribbing, have the apical whorls proportionately much larger, and have the aperture occupying a much larger part of the shell length. The 

points outlined above are only the most obvious set of 

differences; many additional details of structure also 

are different. If unusual species in these families do 

develop features equivalent to those found in Dendropupa, the methods of constructing the characters are 

very distinctive and immediately distinguish those 

species from Dendropupa.

Three families Bulimulidae, Urocoptidae, and 

Enidae require more detailed comparisons with 

Dendropupa. The Neotropical and Austro-Pacific 

Bulimulidae, except such genera as those often 

clumped as Bostryx, have prominent and complex 

apical sculpture. This sculpture consists of both radial 

and spiral lirae, but never, so far as we are aware, incised radial lines. The genera grouped as Bostryx 

(sensu Zilch, 1959-1960) are found mainly in the Andean countries, Chile, Peru, and Bolivia; a few species 

of the subgenus Lissoacme are found in Ecuador and 

Argentina. Specimens of Bostryx have macroscopically smooth apices, do not have the lip expanded or 

reflected, and have irregularly rugose postapical 

sculpture. The whorls are mostly flat sided, the pattern of coiling is very tight (except in Lissoacme), and 

there are no barriers or expanded columellas. The resemblances to Dendropupa are only of a general nature in regard to shape; specific structural features are 

different. No other groups of Bulimulidae, except uncommon"freak" species, have this shape and whorl-increase pattern.

The Urocoptidae are a logical group with which to 

compare Dendropupa. Urocoptids generally have a

radically expanded columella; the columella is armored by one to many spiral ridges, hooks, barriers, or 

corrugations. These features rarely are visible from 

the shell aperture. The shell generally has prominent, 

often sharply defined radial ridges after a smooth or 

weakly striated apex that may have radial ridges on 

the lower parts. Thus the expanded columella, columellar ridges, and radial ribs of Dendropupa suggest 

urocoptid affinities. The choice by Dawson (1880, p. 

408) of Microceramus gossei (Pfeiffer, 1846) for comparison with Dendropupa is not unreasonable. However, Pilsbry (1903-1904, p. xvii, xviii) noted that the 

pattern of axial structures has evolved independently 

several times within the Urocoptidae and also in some 

achatinoid groups. His statement that "homologous 

regions of the shell or its axis have given rise to similar 

structures, wholly independently, in various phyla of 

the family," followed by a list of such convergences 

among genera, points out the improbability of making 

direct comparisons between Dendropupa and extant 

genera of the Urocoptidae.

Although the columellar expansion, ornamentation, 

and shell sculpture are similar in urocoptids and Dendropupa, in several features Dendropupa is quite different from the urocoptid growth pattern. Except in 

such genera as Microceramus and Holospira (sensu 

lato), the spire of urocoptids is normally decollated. 

The apex of the shell is bulbous and smooth to weakly 

ribbed, and the early pattern of spire growth is constricted, commonly leading to concave sides on the 

spire. The whorls are narrow and often exceed 20. 

Even in these genera that do not have a decollated 

spire, the pattern of a bulbous apex having narrow 

whorls and a constricted spire outline persists.

Equally important in differentiating Dendropupa 

from the Urocoptidae are their patterns of apertural 

structure. In nearly all groups of the Urocoptidae, the 

parietal wall either is detached from the penultimate 

whorl, forming a continuous lip around the aperture, 

or has a heavy raised callus. In all these taxa, either 

the palatal lip is simple to weakly expanded, or, if the 

palatal lip is expanded, it is abruptly reflexed and narrow. Only Microceramus and Pineria (Pilsbry, 1903- 

1904, pis. 1, 15, 20-27) depart from this pattern and 

have neither a parietal lip nor a heavy parietal callus. 

Even in these taxa, the lip structure differs greatly 

from that found in Dendropupa. At its upper margin, 

the palatal lip turns abruptly before inserting onto the 

penultimate whorl (Pilsbry, 1903-1904, pi. 26 for example), because the point of attachment is very low on 

the wall. Pilsbry (1946, p. 109) indicated that "Microceramus is a relatively unspecialized member of the 

primitive Antillean Urocoptid stock," but the growth
ANALYSIS AND CLASSIFICATION OF TAXA 13

pattern of the aperture and resultant insertion of the 

palatal lip is such that only lip extension is required to 

produce the detached parietal lip that is characteristic 

of most Urocoptidae.

In contrast, Dendropupa differs in the following 

ways: it does not have a bulbous apex; the spire sides 

are convex; the whorls are few in number (as many as 

nine); the whorls are relatively wide and have curved 

sides; the palatal lip inserts high and does not curve 

before inserting onto the penultimate whorl; no thick 

parietal callus is present; and the palatal lip is broadly 

and gradually expanded (see Dawson, 1880, p. 407, 

figs. 1,2). The Urocoptidae have a basic growth "gestalt" that leads to decollation, detached parietal lip, 

and narrow whorls. Dendropupa does not share this 

basic pattern of shell form and growth. In Dendropupa, the special feature of an expanded columella 

having ridges duplicates a structure characteristic of 

the Urocoptidae, but this is a highly variable feature 

that has evolved several different times. The basic differences in shell form and growth pattern lead us to 

reject the idea that Dendropupa and the Urocoptidae 

are related forms.

The Enidae appear to be the most likely group with 

which to associate Dendropupa. A comparison of the 

illustrations in the latest monograph of the Enidae 

(Kobelt, 1902) with the urocoptid patterns indicates 

that the apertural shape, lip expansion and attachment, apex size, and whorl contours in enids closely 

match the basic growth form in Dendropupa and provide a distinct contrast to those characters typifying 

the Urocoptidae. The basic "gestalt" of shell form in 

the Enidae and Dendropupa is a close match, but few 

enids show major radial sculpture, and most enids 

have a narrow columella. Only in such taxa as Bullminus labrosus labrosus Olivier, 1804 (this report, fig. 

1C) is a ridge developed on the slightly expanded columella. Most enids do not develop a regular radial 

sculpture, but such sculpture has evolved sporadically.

Thus, in basic shell growth form and whorl characteristics, Dendropupa is similar to the extant Enidae; 

in having an expanded columella with two ridges and 

in having heavy radial sculpture, Dendropupa is similar to the extant Urocoptidae and differs from the Enidae. We give greater weight to the basic shell form 

and growth than to the details of the columella and 

shell sculpture. Therefore, we place the Dendropupinae Wenz, 1938, under the Enidae. The features 

that distinguish Dendropupinae from other subfamilies of the Enidae are the expanded columella with 

ridges and the regular radial sculpture on the postapical whorls.

Distributional differences between the extant Urocoptidae and the Enidae are striking. The Urocoptidae has two centers of diversity Mexico and the 

Greater Antilles. Members of both complexes extend 

short distances into North America and on the southern fringes of the Caribbean. Probable urocoptids 

(identified as Holospira) are known from the Eocene 

of both Wyoming and New Mexico and from the Upper Cretaceous of Alberta (Tozer, 1956, p. 82-84), but 

no South American fossil records exist for the Urocoptidae (Parodiz, 1969). The origin in time and space of 

the Urocoptidae is unknown, and its affinities, other 

than that it is loosely associated with the Bulimulidae, 

are unknown. The Enidae is primarily Palearctic and 

has extensive speciation in the Himalayas and Middle 

East and several groups radiating in both tropical Africa and South Africa. A few species are found in Indonesia; a peculiar group, Draparnaudia, is found in 

New Caledonia and the New Hebrides (Solem, 1959; 

1962, p. 219), and one genus is in northern Australia 

(Solem, 1964).

Subfamily DENDROPUPINAE Wenz, 1938

Description.—Columella expanded on lower spire, 

hollow, surmounted by one or more spiral barriers 

that stop well behind the aperture. Apex of shell has 

weak radial incised lines, postapical sculpture of inverted V-shaped radial ribs. Aperture without parietal lip or heavy callus, insertion of palatal lip into 

preceding whorl direct.

Discussion.—The expanded lower columella with 

raised ridges and the strong radial postnuclear sculpture are the main features separating the Dendropupinae from the other subfamilies of the Enidae. We 

recognize only one genus, Dendropupa Owen, 1859, 

with Strophella Dawson, 1859 (=Strophites Dawson, 

1880, not Deshayes, 1832), a subjective synonym.

Genus Dendropupa Owen, 1859

Dendropupa Owen, 1859, p. 94. Type species: Pupa vetusta Dawson, 1859; monotypy. (see Knight, 1941, p. 98) 

Strophites Dawson, 1880, p. 413. Not Strophites Deshayes, 1832. 

Type species: Strophites grandaevus Dawson, 1880; monotypy. 

Strophella Dawson, 1895, p. 84. Type species: Strophites grandaevus Dawson, 1880; monotypy.

Description.—Shell elongate, cylindrical, has about 

nine whorls when adult, spire sides convex. Apical 

sculpture of fine radial lines, postapical of prominent 

narrow radial ribs. Lip expanded, palatal margin inserting directly onto preceding whorl, no parietal lip 

or callus. Lower columella expanded, has raised 

ridges.

Discussion.—The type species is Pupa vetusta 

Dawson, 1859, by monotypy, and it has been discussed
14 NORTH AMERICAN PALEOZOIC LAND SNAILS, SUMMARY OF OTHER PALEOZOIC NONMARINE SNAILS

in detail above. We include Strophites grandaevus 

Dawson, 1880, and Pupa primaeva Matthew, 1895, in 

Dendropupa, because the limited data on these two 

taxa provide no criteria by which they can be separated generically from the type species. In addition, the 

European species D. walchiarum Fischer, 1885, from 

France and D. zarecznyi Panow, 1936, from Poland fit 

quite well within our concept of the genus Dendropupa. The known distribution of Dendropupa thus 

extends from eastern Canada to Poland.

Dendropupa primaeva (Matthew, 1895) 

Figure 2

Pupa primaeva Matthew, 1895, p. 98-99, pi. I, figs. lOa, b; Pilsbry,

1926, p. 318. 

Dendropupa primaeva (Matthew) Henderson, 1935, p. 149; Yen,

1949, p. 236; Yochelson and Saunders, 1967, p. 125.

Despite repeated inquires during a period of several 

years, we have not been able to locate the single specimen on which this species was based nor to find any 

additional material that could be referred to this 

taxon. We have reproduced Matthew's original description (below) and illustrations (this report, fig. 2).

Matthew's (1895, p. 98, 99) description is:

Shell cylindrical, abruptly conical at the apex, increasing rapidly 

in the upper whorls. Whorls 6; sides of the whorls rather flat, upper 

edge sub-carinate; the last whorl about as wide as the shell, somewhat conical below, nucleus very small. Suture impressed. Aperture unknown. Shell surface smooth and shining.

Sculpture.—Surface ornamented with numerous sharp ribs, 

transverse to the whorl, about 20 visible on one-half of a whorl. 

These ribs are not all equally spaced or equally prominent; onesixth of a whorl may have them nearly twice as far apart as the rest. 

While quite distinct on the two last whorls, the ribs are less so than 

on the upper whorls of the shell.

Shell.—Length, 5¥2 mm.; width, nearly 2 mm.

The original illustrations are reproduced here as 

figure 2. The whorl number, whorl contour, shell

Trans. R.S.C., 1894

FIGURE 2. Original illustrations of Dendropupa primaeva 

(Matthew, 1895, pi. I, figs. lOa, b).

shape, size, and sculpture of radial ribs are consistent 

with this species being a small relative of. Dendropupa 

vetusta. We can list no characters that would suggest 

generic separation. As pointed out by Matthew, D. vetusta has more whorls, but otherwise is very similar. 

Tentatively, this taxon is retained as a different species, although the reported diameter of D. primaeva 

(nearly 2 mm) is only marginally below the observed 

range of 2.22-3.27 mm for D. vetusta.

Dendropupa grandaeva (Dawson, 188O) 

Plate 1, figure 4.

Strophites grandaevus Dawson, 1880, p. 413-414, fig. 15; White,

1883, p. 455, 475, pi. 1, fig. 1; Pilsbry, 1926, p. 319, pi. 32, fig. 16;

Henderson, 1935, p. 149. 

Strophella grandaevus (Dawson), Dawson, 1895, p. 84; Wenz,

1938, p. 470, fig. 1195; Knight, 1941, p. 341, fig. 5; Yen, 1949, p.

237. 

Dendropupa grandaevus (Dawson), Yochelson and Saunders,

1967, p. 64.

This species is known only from the holotype, catalogued as number 2384 in the Redpath Museum. It 

has not been restudied since its original description. 

The specimen is preserved apertural side downward 

on a thin slab of shale. This slab has broken so that a 

prominent crack runs diagonally across the lower two 

whorls (pi. 1, fig. 4). A photograph made many years 

ago and attached to the cardboard block holding the 

slab shows the crack as being somewhat narrower than 

it is now. The left margin of the specimen is broken, 

and the right margin is covered by a film of matrix; 

attempts to remove this without injury to the underlying specimen have been unsuccessful. Because neither 

margin of the shell can be seen, Dawson apparently 

mistook the preserved shape for the true form and his 

drawing suggests a bent shell. This interpretation has 

always been one of the enigmatic features of Strophella.

We interpret the holotype as representing a flattened periostracal fragment of a snail shell. There is 

no evidence of any thickness to the fossil. The low relief of the ribs is greater than would be expected, considering the degree of sutural crenulation in areas of 

whorl overlap. Dawson (1880, fig. 15) greatly exaggerated both rib height and degree of sharpness in rib 

contour. His drawing also gives a false impression of 

the shape, suggesting that the specimen had a curved 

columella. The appearance of this fossil is consistent 

with that of a decalcified periostracal shell remnant 

that has been pressed flat. This remnant probably included early postapical to lower spire whorls. The 

body whorl part behind the aperture may be partly 

present.

On the uppermost whorl, the intervals between ribs 

range from 0.12 to 0.14 mm. No accurate measure-
ANALYSIS AND CLASSIFICATION OF TAXA 15

ments can be made on rib spacing on the second 

whorl. On the third whorl, the distances between 

successive parts of ribs generally increase along the direction of growth (right to left); the intervals (right to 

left) are 0.20, 0.26, 0.23, 0.30, 0.33, 0.33, and 0.36 mm. 

On the last whorl, the intervals (right to left) are 0.33, 

0.30, 0.36, 0.40, 0.33, 0.40, 0.36, and 0.40 mm.

Certain predictions concerning shell size can be 

made on the basis of whorl measurements and the intervals between ribs. In modern cylindrical land 

snails, the common pattern of rib spacing is for it to 

increase slightly and regularly as the whorl width increases regularly. If whorl width charges rapidly, rib 

spacing increases dramatically. When maximum shell 

diameter is approached and whorl width is not increasing, the intervals between ribs remain relatively 

constant. On the basis of these tendencies, one may 

infer that Strophella grandaeva may have had a narrow apex, and the early spire may have expanded 

rather rapidly in width. The penultimate whorl of the 

holotype still was widening slightly, as shown by the 

progressive increase in rib interval. On the last whorl 

of the holotype, the basically stabilized rib interval 

suggests that whorl width remained constant. From 

this point of growth, two patterns are equally possible. 

Either shell width stabilized and growth continued for 

a while before a lip formed, or, alternatively, gerontic 

growth and lip formation began almost immediately. 

In either growth pattern, it seems possible that an 

apex of one or two whorls was superimposed above the 

first preserved fragment.

Measurements of individual whorl heights, based 

on rib length from suture to suture, are: top whorl, 

0.93 mm; second whorl, 1.83 mm; third whorl, 2.23 

mm; and last whorl (including basal extension), 3.16 

mm. The shell fragment has a cumulative total length 

of 8.37 mm, although distortions have changed the actual tip to base length slightly. Allowing for a minimum of two low apical whorls, perhaps having a 

combined height of 0.5 mm, plus another half whorl of 

body whorl growth adding 1.5 mm, a total height of 

about 10.4 mm with QVz whorls would be a minimum 

adult size. Growth of two additional adult whorls 

could be possible, and an adult height of as much as 15 

mm is within the realm of speculation.

Clearly, this species falls near the size range of Dendropupa vetusta. Specimens of that species reach a 

height of 12 mm and have as many as nine whorls. The 

apparent major difference in shape is spurious. Indeed, if one allows for the extreme flattening of the 

specimen of S. grandaeva, it and specimens of D. vetusta are similar in profile. The two species do differ 

in the pattern and width of ribbing; Strophella grandaeva shows much more rapid whorl increment.

We judge that generic separation cannot be maintained, and thus we place Strophella as a subjective 

synonym of Dendropupa. Knight, Batten, and Yochelson (1960, p. 1318) had tentatively suggested this 

synonym, but they had not examined the type specimens of either genus.

Anthracopupa ohioensis Whitfield, 1881

Plate 1, figures 1-3, 8-10,13, 14,18, 19; 

plates 5-7; plate 9, figure 6; figures 3-5

Anthracopupa ohioensis Whitfield, 1881, p. 126, figs. 1-4; White, 

1883, p. 456, pi. 3, figs. 5-8; Whitfield, 1891, p. 591, pi. 12, figs. 

15-17; Henderson, 1935, p. 150; Yen, 1949, p. 236; Zilch, 1959- 

1960, p. 63, fig. 202; Yochelson and Saunders, 1967, p. 31.

Anthracopupaf?) dunkardana Stauffer and Schroyer, 1920, p. 

144-145, pi. 11, figs. 12-16; Yochelson and Saunders, 1967, p. 32.

Naticopsis(?) diminuta Stauffer and Schroyer, 1920, p. 145, pi. 11, 

figs. 22-23; Tilton, 1930, p. 112, pi. 4, fig. A; Yochelson and 

Saunders, 1967, p. 140.

We have been able to examine the type of all three 

species considered here under A. ohioensis. The holetype (pi. 1, figs. 13, 14, 18, 19) of A. ohioensis is 

AMNH 8487/1 and is the same morph as the typical 

rather squat form (fig. 4D), although the holotype reduced palatal ridges. The types of A.(?) dunkardana 

are cataloged in the Ohio State University Orton Museum as OSU 15151-15164. The largest and most 

elongated specimen, OSU 15161, is here designated as 

lectotype (pi. 1, figs. 11,12), and the other three specimens are thus lectoparatypes. All examples have juvenile apertures and show no thickening of the lip or 

traces of the palatal barriers. The original figures of 

A.(?) dunkardana were misleading in that the columellar barrier was omitted. The types ofNaticopsis(?) 

diminuta are OSU 15169 and OSU 15170. They are 

very young shells, still partly imbedded in matrix. The 

larger specimen, OSU 15170, is here designated lectotype (pi. 1, figs. 1, 2). It has a very wide apical angle 

and would have grown into one of the very short and 

broad adults, like that shown in figure 4D.

In addition to these types, we have about 100 specimens in the FMNH that were collected by P. D. 

Moore and associates and 500 specimens in the USGS 

collections (Yochelson, 1975).

Inclusion in one species of the diverse-appearing 

morphotypes represented by the three names and the 

examples in figures 3-5 requires justification. Because 

of the abundant material now available, variation 

both within individual collections (table 2) and in the 

pooled sample could be analyzed. Approximately 600 

specimens were studied, of which 230 were adult and 

were sufficiently undistorted and complete to allow at 

least one measurement to be made. Shell height, shell
16 NORTH AMERICAN PALEOZOIC LAND SNAILS, SUMMARY OF OTHER PALEOZOIC NONMARINE SNAILS

diameter, and whorl count could be measured accurately in many specimens. The height/diameter ratio 

was calculated where possible. Because of flattening 

during preservation, broken edges on the whorls and 

spire, apical wear, loss of lip parts, and similar problems, each measurement could not be made on every 

specimen. When distortions or broken pieces prevented an accurate measurement, that measurement was 

not made on the specimen. For 226 specimens, shell 

height could be recorded; for 187 specimens, the whorl 

count was determined to the nearest eighth whorl; for 

182 specimens, the shell diameter was measured; and 

for 179 specimens (of which both height and diameter 

had been measured), the height/diameter ratio was 

calculated. Partial crushing of the shell body whorl or 

general flattening during preservation was quite common, so that shell diameter commonly could not be 

determined. The remaining several hundred specimens were fragmentary, juvenile, imbedded in matrix, 

or too distorted to be measured. Table 2 lists the data 

of specimens from localities that have yielded more 

than a few scattered specimens well enough preserved 

to be measured.

The fossil localities are spread over a small geographic area relative to the range of many modern 

species and include deposits of slightly different ages. 

All samples contain adults that are very similar in size 

and shape. Mean whorl counts listed on table 2 are essentially identical, and the samples show an equally 

wide variation. Shells from the Nineveh Limestone 

Member, Greene Formation, were on the average a little smaller in diameter. A sample from the "Lower 

Washington Limestone Member," Washington Formation, lacked the more elongated morph. The differences among samples, particularly in view of the large 

ranges in variations, are not significant. When the 

samples are pooled (table 3), the continuous nature of 

variation is evident. The height and whorl-count distributions, which are closely linked in expression, 

have an extended mode because of the variability, but 

no indication of any bimodality exists. Similarly, table 

4, showing the relation of shell height to shell diameter, and table 5, showing the relation of shell height to 

whorl count, demonstrate normal distributions without any trace of bimodality. On the basis of size and 

shape variation, we cannot distinguish any subgroups 

within A. ohioensis.

Study of thin sections prepared from material in 

both the Little Captina Limestone Member and 

"Lower Washington Limestone Member" confirmed 

the interpretation of A. ohioensis shell structure inferred from the SEM photographs. In the cross-sectional view (fig. 3), one can see a very thin periostracal 

layer between the calcium layers of the shell; the existence of the periostracum indicates that it was not resorbed prior to the attachment of the lower whorls 

(see pi. 5, figs. 2-4 for the gap that exists where the 

periostracum was removed before preservation). The 

early whorls are not resorbed, a variable umbilical 

chink exists in the columella (ruled areas in fig. 3) during stages of growth, and the barriers do not extend up 

into the inner whorls.

The variability of A. ohioensis involves not only size 

and shape, but also the degree of development of both 

lip reflection and apertural barriers. Figure 4 shows 

four selected stages of barrier development. These

TABLE 2 Size (in mm) and shape variation in samples of Anthracopupa ohioensis Whitfield, 1881

[N, number of specimens. SEM, standard error of the mean]

Stratigraphic units 

(collection nos.)

"Lower Washington" 

(24846,24855-7) _ __

Nineveh 

(24827)

Monongahela (Uniontown) 

(24851 2 collections) _ _

Little Captina 

(24854)

(24840)

(24836)

(24850)

Scott's Cut (FMNH) ___

N

18 

34

16 

44

15

21

12

47

Height 

Mean ± SEM 

(range)

2.92 ±0.036 

(2.66-3.13)

3.03 + 0.076

(2.66-4.14)

3.15 + 0.082 

(2.76-4.01)

3.24 + 0.036

(2.73-3.78) 

3.29 + 0.078

(2.86-3.91) 

3.17 + 0.075

(2.66-3.75) 

3.21 + 0.070

(2.76-3.59) 

3.15 + 0.031 

(2.80-3.59)

N

9 

31

13 

23

16

15

11

47

Diameter 

Mean ± SEM 

(range)

1.82 ±0.033 

(1.71-2.01)

1.72 + 0.042

(1.35-2.17)

2.04 + 0.030

(1.84-2.17)

1.88 + 0.026

(1.61-2.11) 

2.02 + 0.031

(1.84-2.17) 

1.86 + 0.030

(1.71-2.04) 

1.99 + 0.038

(1.78-2.17) 

1.94 ±0.015 

(1.71-2.14)

N

9

30

13 

23

14

15

11

47

Height/diameter 

ratio 

Mean ± SEM 

(range)

1.62 ±0.049 

(1.38-1.83)

1.75 + 0.031

(1.39-2.09)

1.57 ±0.042 

(1.31-1.85)

1.71 + 0.033

(1.45-2.02) 

1.65 + 0.044

(1.49-1.95) 

1.71 + 0.037

(1.46-2.00) 

1.60 + 0.045

(1.45-1.98) 

1.63 ±0.015 

(1.42-1.88)

N

8 

32

13

17

14

19

11

38

Whorl Count 

Mean ± SEM 

(range)

4.83 ±0.071

(4i/2 _5Vs)

4.81 + 0.047 (4'/2-5y2)

4.76 ±0.046

(4 I/2-5'/fe)

4.90 + 0.049

(4 1/2 -5%) 

4.83 + 0.075

(4 1/2-5 I/4)

4.86 + 0.051

(4 1/2-5 1/4)

4.94 + 0.052

(4 3/4-5'/4)

4.79 ±0.033

(4 1/2-5'/4)
ANALYSIS AND CLASSIFICATION OF TAXA 17

TABLE 3. Frequency distribution for shell height, shell diameter, height/diameter, ratio, and whorl count in pooled samples of adult

materials of Anthracopupa ohioensis Whitfield, 1881

Shell height (mm)

Range

2.53-2.66

2.67-2.83

2.84-2.99

3.00-3.16

3.17-3.32

3.33-3.49

3.50-3.65

3.66-3.82

3.33-3.98

3.99-4.14

No. of 

specimens

4

34

49

49

35

30

10

5

5

5

Shell diameter (mm)

Range

1.35-1.48

1.49-1.64

1.65-1.81

1.82-1.97

1.98-2.14

2.15-2.30

_ _

_ _

_ _

--

No. of 

specimens

5

15

35

77

39

11

_ _

_ _

_ _

--

Height/diameter ratio

Range

1.30-1.39

1.40-1.49

1.50-1.59

1.60-1.69

1.70-1.79

1.80-1.89

1.90-1.99

2.00-2.09

_ _

--

No. of 

specimens

3

15

47

47

31

20

10

5

_ _

--

Whorl count

Mean

4%

4 5/8

4%

4%

5

5 '/s

5V4

5%

5V2

--

No. of 

specimens

24

27

36

36

31

20

10

3

1

--

TABLE 4. Relation of shell height to shell diameter in all adult specimens of Anthracopupa ohioensis Whitfield, 1881

Shell height (mm)

o KQ o 70

2.71-2.89

2.90-3.09

3.10-3.29

3.30-3.49

3.50-3.68

3.69-3.88

3.89-4.08

Total

Shell diameter (mm)

1.35- 

1.41

1

__

__

__

__

1

1.42- 

1.51

1 

4 

3

8

1.52- 

1.61

1

5 

2

8

1.62- 

1.71

1

7 

2 

2 

1

13

1.72- 

1.81

9

5 

7 

2 

2

25

1.82- 

1.91

10

17 

7 

5 

1

1

41

1.92- 

2.01

3 

13 

11 

10 

3 

2 

1

43

2.02- 

2.11

2 

5 

10

5 

1 

1 

2

26

2.12- 

2.20

2 

5 

2

4

13

2.21- 

2.30

1

1

Total

3 

41 

49 

42 

25 

8 

4 

7

179

TABLE 5. Relation of shell height to whorl count in all adult specimens of Anthracopupa ohioensis Whitfield, 1881

Whorl count

Shell height (mm)

2.53-2.70

2.71-2.89

2.90-3.09

3.10-3.29

3.30-3.49

3.50-3.68

3.69-3.88

3.89-4.08

4.09-4.27

Total __________

4'j

2

10

8

4

__

__

__

--

24

4-'-

2

9

10

4

2

__

- _

27

4'4

__

12

11

8

3

1

__

--

35

4 7 H

__

3

11

13

6

1

1

- -

35

5

__

3

5

8

12

3

31

5' 8

__

1

1

7

6

4

1

20

5'4

_ _

__

_ _

1

1

1

2

4

1

10

5 J » 5"2

_ _ _ _

__

_ _ _ _

_ _ _ _

_ _ _ _

2

1 1

3 1

Total

4

38

46

45

30

10

5

7

1

186

features and those shown in figure 5 and on plate 5, 

figures L-3, demonstrate that shell shape and apertural features are not correlated. Even though many of 

the smaller squatter shells do have huge teeth and 

broadly reflected lips (figs. 4C, D), the same lip features and huge teeth occur in elongated slender shells 

(pi. 5, fig. 1). The development of the upper palatal 

ridge takes place very late in ontogeny, as discussed 

below, and the ridge may be absent from specimens 

having thin lips (fig. 4B). The lower palatal ridge varies from wide and bifurcated (pi. 5, fig. 2; figs. 4C, D) 

to narrow and single (fig. 4A). The inward sinuation of 

the lower palatal lip varies from absent (figs. 4A, B) 

through slight (pi. 5, fig. 1) to very prominent (fig. 4C).

Variation in thickness of the lip does correlate with 

the degree of palatal ridge development. If the lip is 

very thick and has grown inward, the palatal ridges 

are large. If the lip is thin and only slightly expanded, 

the palatal ridges are absent or only weakly developed. After initial reflection of the lip (fig. 5A), the 

subsequent growth is internal, narrowing the aperture 

by adding great thickness to the palatal wall and then 

building the palatal ridges. The lip thickening and palatal-ridge development are ontogenetic changes (figs. 

5A-C).

Despite attempts on our part, we could find no basis 

on which to divide this material into two or more taxa. 

Anthracopupa ohioensis appears to be a highly vari-
18 NORTH AMERICAN PALEOZOIC LAND SNAILS, SUMMARY OF OTHER PALEOZOIC NONMARINE SNAILS

2mm

FIGURE 3. Polished section of Anthracopupa ohioensis Whitfield, 1881. Little Captina Limestone Member, Monongahela Formation, USGS locality USGS 

24840-PC (USNM 242, 697). Ruled areas indicate 

hollow parts of columella. Periostracum is continued 

between new whorl attachment sections (see pi. 5, 

figs. 2-4, for an example of a specimen from which the 

periostracum has dissolved away). Note retention of 

the upper whorl partitions and slight remnant of basal ridge in section.

able species, as noted above, but it is not bimodal in its 

variation. Whenever more than 5 or 10 specimens 

were available from a locality, the same range in variation was found. We conclude that only one species 

can be recognized.

Preservation of the material varies greatly. The material includes badly crushed fragmentary specimens, 

steinkerns of the inside whorl profile, and recrystallized exact replicas of the shell surface. In one sample,

from the "Lower Washington Limestone Member" in 

Ayers limestone quarry (USGS 24855-PC), a miracle 

of preservation permitted us to study shell details at 

nearly 10,000 X (pi. 7, figs. 1-4). The following observations on structure were made about a variety of 

specimens and preservation types.

Commonly, specimens preserved in various ways 

were found at a single locality. Illustrated specimens 

from USGS 24855-PC indicate two extremes. These 

are easily separated visually by the presence (pi. 6, 

figs. 1-3) or absence (pi. 5, fig. 1) of radial sculpture on 

the spire. Specimens without the radial sculpture 

showed different layers of crystals in what was originally the calcareous part of the shell (pi. 5, figs. 4-6) 

and no microprojections on the surface of the major 

barriers. Specimens having radial ribbing preserved 

showed only one type of crystallization (equivalent to 

that seen in Dendropupa uetusta, pi. 4, figs. 4-5) and 

many microprojections on the surface of the barriers 

(pi. 7, figs. 1-4). We interpret the mode of preservation of the specimens not having radial sculpture as 

loss of the periostracum without its replacement, followed by surface erosion of the barriers. We interpret 

the second type of preservation as replacement of 

both periostracum and calcareous layers by a single 

crystalline structure; exquisite surface details were 

preserved in the process. We have not studied the frequency of these two types of preservation or the mineralogy of the replacing mineral.

Examination of a ribless adult shell having well-developed barriers (pi. 5) shows that the parietal and columellar barriers (pi. 5, figs. 2-4) are relatively simple 

in structure and position and normally show no 

change from the locations and directions present in 

the juveniles (pi. 6, figs. 3-6). A speciment having a 

greatly distorted lip (figs. 4C, D) may also have the 

columellar barrier angled slightly upwards. In contrast, the palatal ridges are totally absent in juveniles 

(pi. 6, figs. 3-6), start to appear in specimens having 

newly formed lips (figs. 4B, 5A) and are prominent 

only in gerontic shells (pi. 5, figs. 2, 3; figs. 4C, D). The 

lower palatal ridge is basal in position and normally is 

quite broad and has only a slight sinuation on its upper surface. The outer palatal ridge (pi. 5, fig. 3) commonly consists of two knobs connected by a lower, but 

still elevated, ridge.

The parietal and columellar barriers of the ribbed 

juvenile shells (pi. 6, fig. 1) are narrower than those of 

the gerontic individuals, and the columellar barrier 

shows a slightly curved tip (pi. 6, figs. 4, 6). The microstructure of the barriers is preserved to differing 

degrees. The surface radial sculpture on the shell spire 

is missing from the gerontic individual illustrated on 

plate 5, and apparently the surface layers of the bar-
ANALYSIS AND CLASSIFICATION OF TAXA 19

2mm

FIGURE 4. Range of variation in barrier development of Anthracopupa ohioensis Whitfield, 1881. "Lower Washington Limestone 

Member," Washington Formation. A, USGS 24857-PC. B, USGS 24855-PC. C, D, USGS 24846-PC.

riers also are absent (pi. 5, figs. 4-6). On plate 5, figure

4. a crevice is discernible between the end of the parietal barrier with its continuing callus and the surface 

of the penultimate whorl. We interpret this crevice as 

representing a missing periostracal layer that had 

been dissolved after death and burial but before diagenetic modification of the shell layers had taken 

place. More detailed examination of the barrier edge 

(pi. 5, figs. 5-6) shows two different crystalline layers, 

an inner layer (left of pi. 5, fig. 5) consisting of large 

horizontal crystals, and an outer layer of vertical columnar crystals (pi. 5, figs. 5-6). This outer layer was 

covered by a second complex layer (see right part of pi.

5. fig. 5), details of which could not be studied because 

of surface wear. The photograph fracture section was 

chipped during cleaning; hence, the two crystal layers 

were unworn.

Several specimens from Ayers quarry had the periostracal layers apparently intact at the time of preservation. Study of details on these specimens (see also 

fig. 3) produced critically important data in determining the affinities of Anthracopupa. The apical whorls 

are smooth, even at 205X magnification (pi. 6, figs. 

1,2), whereas the postapical whorls (pi. 6, figs. 2, 3, 5) 

have a sculpture of low, rather closely spaced axial 

ribs. Remnants of this form of sculpture also were 

seen on the juvenile Anthracopupa that was part of 

the type lot of Pupa bigsbii (pi. 8, fig. 3, upper left 

side). The ribs are greatly reduced below the periphery of the whorls (pi. 6, figs. 3, 5), and this perhaps

explains the existence of the crevice mentioned above 

and shown on plate 5, figure 4. If the rib profile is very 

low or reduced below the periphery, resorption of the 

periostracal covering and (or) ribs is not necessary before growth of the next whorl. This contrasts with the 

situation in extant endodontoids, where rib and periostracal resorption on the lower part of the whorl is 

necessary preliminary to growth of the next whorl (Solem, 1973a, fig. 7). Preservation of these Anthracopupa seems to have involved replacement of both 

shell and periostracum.

The barriers in the specimens having preserved periostracum showed incredibly minute surface detail 

(pi. 7). An excised parietal barrier was mounted for 

SEM observation (pi. 7, fig. 1) in order to minimize 

electron charging interference. Matrix in the aperture 

(pi. 6, figs. 3, 4, 6) charged badly, and higher magnification study of the parietal barrier would have been 

impossible without separating it from the matrix. The 

sides of the barrier have very few microprojections (pi. 

7, fig. 2), and the top of the barrier is crowded with 

very large microprojections (pi. 7, figs. 3, 4); this pattern is neither accidental nor an artifact. In some recent Pupillidae, such as Vertigo morsel Sterki, 1894 

(Solem and Lebryk, 1976, figs. 27-32), exactly the 

same pattern is seen; microprojections on the barrier 

tops are much larger than those on the lip edge and 

sides of the barriers. As is typical in most pupillids, 

the microprojections are roughly triangular (pi. 7, fig. 

4). Even on a shell embedded in matrix (pi. 7, figs. 5,
20 NORTH AMERICAN PALEOZOIC LAND SNAILS, SUMMARY OF OTHER PALEOZOIC NONMARINE SNAILS

2mm

FIGURE 5. Ontogenetic development of expanded lip 

and palatal ridges in Anthracopupa ohioensis 

Whitfield, 1881. A, Newly adult. B, Typical adult. 

C, Gerontic individual. A, B, From Monongahela 

Formation, Scott's Cut, Warnock, Ohio (FMNH). 

C, From "Lower Washington Limestone Member," 

Washington Formation, Ayers limestone quarry, 

Belmont County, Ohio (USGS 24855-PC).

6), the form of the barrier microprojections remains 

relatively unworn and is suitable for study after very 

careful cleaning.

As is discussed later in this report, the data presented above (p. 62-73) are sufficient to classify Anthracopupa ohioensis with the Tornatellinidae; to define the subfamily Anthracopupinae Wenz, 1938; and 

to include several less well known taxa in the genus 

Anthracopupa.

Subfamily ANTHRACOPUPINAE Wenz, 1938

Description. Parietal and columellar barriers typical of family, lip thickening internally and developing 

two ridges only in late growth stages. Apex smooth, 

postapical sculpture of low to strong radial ribs.

Discussion.—Because of the unusual pattern to formation of the shell lip, the great time gap between 

Anthracopupa and the modern Tornatellinidae, and 

the high degree of subfamily and tribal divergence

among the modern taxa (Cooke and Kondo, 1960, p. 

16-18, 50, 51), we accept subfamily-level divergence 

for the Anthracopupinae.

Only one genus, Anthracopupa Whitfield, 1881, is 

recognized and Maturipupa Pilsbry, 1926, is listed as 

a subjective synonym.

Genus Anthracopupa Whitfield, 1881

Anthracopupa Whitfield, 1881, p. 126. Type species: Anthracopupa ohioensis Whitfield, 1881; original designation.

Maturipupa Pilsbry, 1926, p. 317. Type species: Pupa vermilionensis Bradley, 1872; original designation.

Pilsbry (1926, p. 317) separated Maturipupa from 

Anthracopupa as it was originally illustrated on the 

basis of Maturipupa''s much more elongated shell, oblique parietal margin, and shortly ovate aperture. Because these differences represent the extremes of the 

shape and ontogenetic variation within a species as 

discussed above, Maturipupa is here placed in subjective synonymy. The comment of Pilsbry (1926, p. 317) 

that the shell "*** has more resemblance to the Tornatellinidae or to some Auriculidae ( = Ellobiidae) 

than to any Pupillidae, and it probably does not belong to the latter family" was not accepted by subsequent workers. Henderson (1935, p. 150,151) and Yen 

(1949, p. 236) continued to place Maturipupa in the 

Pupillidae; Wenz (1938, p. 52, 68) and Zilch (1959- 

1960, p. 63, 64) grouped the two taxa as a subfamily of 

the Ellobiidae; Knight, Batten, and Yochelson (1960, 

p. 1318) placed them in the prosobranch group Cyclophoridae.

The question of potential family affinities for Anthracopupa is obviously controversial. On the basis of 

the structural features described above, we make the 

following arguments. Reference to the Cyclophoridae, 

or any other prosobranch group, is improbable because Anthracopupa has both a parietal and a columellar barrier. As noted under the discussion of 

Dendropupa, coexistence of major apertural barriers 

and an operculum (other than a remnant) is not possible. Barriers equivalent to those in the endodontid 

land snails are found in only one land prosobranch 

group, the helicinid subfamily Proserpininae. The 

barriers are particularly strongly developed in the 

genera Proserpina Gray, 1839, from Jamaica and 

Ceres Gray, 1856, from Mexico. Prosperinella Bland, 

1865, and Cyane H. Adams, 1870, have reduced barriers. None of these genera have an operculum, and in 

their pattern of shell form and anatomical structure, 

they are clearly helicinid. Anthracopupa and all extant cyclophorid groups do not have features with detailed similarities. The presence of the two barriers 

throughout ontogeny in Anthracopupa led us to exclude prosobranch affinities from further consider-
ANALYSIS AND CLASSIFICATION OF TAXA 21

ation. We cannot exclude the slight possibility that 

Anthracopupa was a nonoperculated prosobranch, 

but because of the striking detailed similarities of 

Anthracopupa to extant pulmonate families, we consider that it was a pulmonate land snail.

The assignment of Anthracopupa by Wenz (1938) 

and Zilch (1959-1960) to the only terrestrial group of 

the superorder Basommatophora, the Ellobiidae, follows a suggestion by Pilsbry (1926); their assignment 

is based in part on errors in the original illustrations. 

Both Whitfield (1881) and Bradley (1872) showed the 

parietal margin as being weakly arched and obliquely 

angled to the shell axis. The typical pupillid-tornatellinid has a much more strongly arcuate and angled 

margin. Illustrations presented here (figs. 3-6) show 

that the parietal margin is well within the pupillidtornatellinid range, and that it is quite different from 

the ellobiid slanted parietal wall (see Zilch, 1959- 

1960, p. 64-78). A general difference also exists in the 

barrier position and structure. In the ellobiids, the parietal barrier, if single, invariably is located near the 

parietal-collumellar margin, and the columella is sinuately twisted and has a ridged barrier. In Anthracopupa (pi. 6, figs. 3-6), the parietal barrier is higher on 

the parietal wall, and the columellar barrier is lamellar and riot sinuate.

The greatest similarity of Anthracopupa to the Ellobiidae is in the method of palatal lip thickening. In 

the Holocene Ellobiidae, such as Pythia Roding, 1798, 

Cassidula Ferussac, 1821, andEllobium Roding, 1798, 

the palatal lip first reflects slightly, then builds up internally and narrows the aperture by inward growth, 

just as in Anthracopupa (fig. 5). We consider this 

similarity to be convergent in nature. The basic shell 

form of the Ellobiidae, even in Carychium Miiller, 

1774 (see Pilsbry, 1948, p. 1051-1061), retains the oblique parietal wall and rarely adopts a pupilliform or 

tornatelliniform shape. Moreover, the shell structures 

of the Ellobiidae and of the higher pulmonates are 

fundamentally different. In the Ellobiidae, the shell 

axis and whorl partitions in the upper spire are resorbed by the animal; they are not resorbed in the pupillid-tornatellinid lineages. The elegant study on 

Pythia by Harry (1951, figs. 9-10) and the illustrations of Carychium in Pilsbry (1948, figs. 562A, 566A, 

567B) show quite well that resorption has taken place. 

Anthracopupa (fig. 3) retains the upper whorl partitions, and thus it cannot be grouped with the Ellobiidae. The lip formations of Anthracopupa and the 

Ellobiidae are similar only very late in ontogeny. The 

differences in parietal barrier positon, parietal wall 

angle, upper whorl partitions, and basic shape are ontogenetically earlier and combine to separate the two 

groups clearly.

The problem then becomes to compare Anthracopupa with extant groups of the superorder Stylommatophora. The shells of groups such as the Enidae, 

Subulinidae, Urocoptidae, Achatinidae, and Bulimulidae, with which Dendropupa was compared, are 

much larger, have more whorls, and have quite different growth patterns than shells of Anthracopupa. The 

zonitidendodontid-helicoid taxa differ in sculpture, 

form, whorl count, and many other features. We consider that small size, presence of apertural barriers, 

and relatively few whorls, very narrow columella (fig. 

3), elongated shell form, and simple shell sculpture, 

narrow the choice down to the pupilloid-tornatellinoid lineages. Both of these groups traditionally have 

been cited as potential relatives of Anthracopupa. We 

must first indicate whether constant shell differences 

exist between the pupilloids and tornatellinoids, and 

then indicate to which pattern Anthracopupa conforms.

The growth pattern shown by the apex and spire is a 

major and characteristic difference between pupilloid 

and tornatellinoid taxa. In the pupilloid groups (fig. 

6A), the upper spire has a distinctly convex outline resulting from significant decoiling. In contrast, the 

typical tornatellinoid form (fig. 6B) results in a nearly 

straight-sided spire outline, or at most (fig. 6C), a very 

slight convexity. These differences hold for almost all 

taxa but are subject to certain growth modifications 

that are easily recognizable as exceptions. To test the 

significance and stability of this difference, all species 

of pupillids illustrated in the comprehensive monograph by Pilsbry (1916-1935) and all the tornatellmids illustrated by Pilsbry and Cooke (1914-1916), 

Odhner (1922), and Cooke and Kondo (1960) were 

scored by Barbara Walden as to whether the spire outline was pupilloid, tornatellinoid, or intermediate. 

Not all named species are illustrated in these papers, 

and no attempt was made to cover pupillids described 

since 1935. Only a few proved difficult to assign to a 

category, as can be seen from the results summarized 

below:

Number of taxa having basic spire form 

(percentages in parentheses):

Pupillids (all) _____ 

Pupillids (normal) _ _ 

Tornatellinids _ _ _ _

Pupilloid

339 (62) 

252 (86) 

0

Intermediate

145 (27)

31 (11)

3( 1)

Tornatellinoid

63 (11)

8( 3)

227 (99)

Two species of tornatellinids having an intermediate 

spire shape have a very blunt apex and expanded early 

spire. They are Tornatellides oncospira Pilsbry and 

Cooke, 1915 (Pilsbry and Cooke, 1914-1916, p. 214, pi. 

46, fig. 8), and T. oswaldi Cooke and Kondo (1960, p. 

257, fig. 109a). The third species, Fernandezia cylin-
22 NORTH AMERICAN PALEOZOIC LAND SNAILS, SUMMARY OF OTHER PALEOZOIC NONMARINE SNAILS

2mm 2mm

FIGURE 6. Shell-form differences in the pupillid (A, D) and tornatellinid (B, C) taxa. Apertural barriers omitted from all these figures. A, Vertigo modesta (Say, 1824) modified from Pilsbry 

(1948, p. 983, fig. 527); B, Lamellidea (L.) oblonga (Pease, 1864) 

modified from Cooke and Kondo (1960, p. 203, fig. 89e); C, Antonella trochlearis fusiforma Cooke and Kondo, 1960 (modified 

from Cooke and Kondo, 1960, p. 121, fig. 54c); D, Bensonella lamothei (Bavay and Dautzenberg, 1912) modified from Pilsbry 

(1916-18, pi. 35, fig. 3).

drella Odhner (1922, p. 242, pi. 9, fig. 49), has a subuliniform shape and a greatly increased whorl count. It 

is very atypical of its genus, which is composed mostly 

of bulimuliform species.

The pupilloid taxa having tornatelliniform spire 

shapes have achieved this shape either by whorl-count 

increase with shell elongation or by becoming squatly 

triangular in outline (fig. 6D). Comparatively few of 

the 50 full pupillid genera are involved in these alterations. A summary of the spire-outline distribution in 

the elongated taxa follows:

Pupilloid

Fauxulus Schaufuss, 1869 ______ 4 

Abida Leach, 1830 __________ 5 

Granopupa O. Boettger, 1889 _ _ _ _ 2 

Chondrina Reichenback, 1828 _ _ _ 3 

Lauria Gray, 1840 __ ______ 37 

Pupoides L. Pfeiffer, 1854 4

Pupisoma Stoliczka, 1870 ______ -- 

Nesopupa Pilsbry, 1900 _______ 32

Totals 87

Intermediate

2 

10 

7 

16 

15 

19

22 

23

114

Number of species having basic spire form (dash leaders indicate 

none found):

Tornatellinoid

4

4

3

7

20

An additional 35 of the tornatelliniform pupillid 

species are found in the squatly triangular genera Gibbulina Beck, 1837, Hypselostoma Benson, 1856, Boysidia Ancey, 1881, Gyliotrachela Tomlin, 1930, 

Systenostoma Bavay and Dautzenberg, 1909, Anauchen Pilsbry, 1917, Aulacospira Moellendorff, 1890, 

and Odontocyclas Schliiter, 1838. Such recently described or separated genera as Ulpia Scott, 1955, Afriboysidia Zilch, 1939, and Bensonella Pilsbry and 

Vanatta, 1900, have the same growth form. The shells 

are essentially triangular in outline and have the 

whorls decoiling regularly down to, and often strikingly on, the body whorl (fig. 6D). In these taxa, the 

whorl count is low, the altered decoiling pattern has 

produced convergence in the spire shape.

Thus, 55 of 63 "tornatelliniform" pupillid species 

are in genera that have greatly altered whorl counts or 

atypical decoiling patterns, and 114 of the 143 pupillids having intermediate shapes are in the few elongated genera that have higher whorl counts. If the 

"typical" or "normal" pupillids are compared with the 

tornatellinids, then the differences in shape are striking and highly stable.

In whorl count and basic shell shape, Anthracopupa is very similar to typical pupillids and tornatellinids, but in spire shape (pi. 5, fig. 1; pi. 6, figs. 1, 3, 5; 

pi. 8, figs. 3, 4; figs. 3, 4), Anthracopupa has the characteristic appearance of the Tornatellinidae.

In yet another fundamental feature, the pupillids 

and the tornatellinids differ without overlap. Tornatellinids, except Achatinella and its Hawaiian relatives, have a few too many apertural barriers present 

in juveniles. These barriers continue in many taxa until or even through the gerontic growth stage. Fully 

adult specimens may have the barriers reduced or absent, but their presence in juveniles is characteristic of 

the family. In contrast, the pupillids only develop the 

apertural barriers when nearly adult or as part of the 

lip-formation syndrome that signals the end of shell 

growth. Although many pupillids are without these 

barriers and have only a reflected lip, the barriers 

themselves are never present in juvenile shells. Young 

Anthracopupa (pi. 6, figs. 3-6) have a well-developed
ANALYSIS AND CLASSIFICATION OF TAXA 23

columellar barrier and a parietal barrier, both of 

which continue into the adult growth stage (pi. 5, figs. 

1, 2; figs. 4, 5), so that the tornatellinid barrier growth 

pattern is followed. The formation of a reflected lip in 

Anthracopupa is comparable with the lip formation 

in Auriculella Pfeiffer, 1855, and in a few Achatinella 

Swainson, 1828; lip expansion is a somewhat unusual 

feature in the Tornatellinidae.

Both tornatellinids and pupillids have microprojections on the barriers (Solem, 1972; Solem and Lebryk, 

1976), but whether or not there are family-level differences in structures of these microprojections cannot 

be determined at this time.

To summarize the above discussion, shell shape, ontogeny of the barriers, and retention of internal early 

whorl sides by Anthracopupa clearly indicate that it 

is not closely related to the Ellobiidae and pupillid 

taxa. The features of Anthracopupa and the Tornatellinidae are compatible; only the unusual lip formation and palatal ridges of Anthracopupa depart from 

the standard tornatellinid pattern. Even the variation 

in shell shape seen in Anthracopupa ohoiensis (figs. 4, 

5) finds a modern counterpart; species such as Lamellidea pusilla (Gould, 1847) see Cooke and Kondo, 

1960, fig. 82a-p) and Tornatellides oblongus (Anton, 

1839) (see Cooke and Kondo, 1960, fig. 105a-l) showing intra- and intercolony variations in shell shape 

and whorl shape.

In addition to Anthracopupa ohioensis, undescribed species of the genus are from near Prague and 

Vienna; A. britannica Cox, 1926, and A. rothenburgensis Hintze, 1933, are from Europe; and three taxa 

are from the Eastern United States 1A. parva 

(Stauffer and Schroyer, 1920), A. sturgeoni Solem and 

Yochelson, n. sp., and A. vermilionensis (Bradley, 

1872). These species differ in details of shape, barrier 

development, size, and lip expansion. No key to identification of species is given because we lack sufficient 

information on the European taxa to permit assessment of their affinities. Comparative comments are 

included under the respective species.

Anthracopupa sturgeoni Solem and Yochelson, new species

Plate 9, figures 1-5)

Diagnosis.—Shell small, slender, heavily ribbed. 

Parietal barrier thick and has a bulbous tip, columellar barrier deeply recessed and reduced in size, lip 

ridges as in A. ohioensis. A. ohioensis less elongated, 

barriers less thickened and modified even on gerontic 

shells, ribbing distinctly weaker and lip normally less 

thickened.

Description.—Shell narrow, elongated, probably 

has 5 3/4 to 6 :/2 whorls, spire not expanded rapidly, last 

whorls not bulbous. Sculpture of apical whorls unknown, lower whorls have prominent, slightly protractive radial ribs, whose interstices are three to four 

times their width. Sutures deep, whorls flatly rounded 

laterally. Aperture ovate, no parietal callus, ribs extend into aperture without modification. Parietal barrier very wide, thickened, has clearly bulbous tip. 

Columellar barrier deeply recessed, reduced in height 

and widened. Basal ridge weakly bifurcated, present 

only in gerontic individuals. Palatal ridge somewhat 

recessed, higher, and slightly bulbous compared with 

basal ridge. Height of shell an estimated 4-4.5 mm, 

diameter 1.31-1.41 mm.

Holotype.—Little Captina Limestone Member, 

Monongahela Formation, at USGS locality, USGS 

24836-PC (USNM 242,705).

Paratypes.—USGS locality 24836-PC, 1 specimen 

(USNM 242,706); USGS locality 24840-PC, 3 specimens (USNM 242,707; 242,708; and 242,709); USGS 

locality 24851-PC, 4 speciments (USNM 242,710; 

242,711; 242,712; and 242,713); USGS locality 24854- 

PC, 1 specimen (USNM 242,714).

Discussion.—All known examples are from the Little Captina Limestone Member of the Monongahela 

Formation, except the specimens from USGS 24851- 

PC, which are from the overlying "Uniontown Limestone Member," Uniontown Formation. No complete, 

undamaged specimens are known. The holotype is the 

nearest to being complete, but it is crushed laterally 

and has the upper spire broken off and twisted to one 

side. Nevertheless, this specimen permits grouping as 

one species the few spire and body whorl fragments 

that constitute the remaining material. Four apical 

fragments lack all sculpture (USNM 242,710- 

242,713), a small midspire fragment has well-preserved sculpture (USNM 242,714), an upper spire has 

good scuplture except on the apex itself (pi. 9, fig. 4, 

USNM 242,706), and three adult body whorl fragments (USNM 242,707-242,709) vary in sculpture and 

in age. In one body whorl fragment without sculpture, 

the lip is only slightly expanded and lacks the ridges, 

the parietal barrier is thinner and without the bulbous 

tip, and the columellar barrier reaches to the lip edge. 

A second specimen from the same locality also is without sculpture, but the lip is slightly thicker and has 

bare traces of the ridges, and the barriers are somewhat thicker. The third specimen also is barely adult, 

and sculpture has been worn off the shell.

The apex and spire (pi. 9, fig. 4) are narrow and have 

flatly rounded whorls and rather deep sutures. A. 

sturgeoni has very slight decoiling, and its prominent 

ribbing contrasts with that of A. ohioensis. The ribs in 

A. sturgeoni are narrow, high, and rather widely 

spaced (pi. 9, fig. 5) compared with those of A. ohioensis (pi. 9, fig. 6). The ribs of A. sturgeoni are more like
24 NORTH AMERICAN PALEOZOIC LAND SNAILS, SUMMARY OF OTHER PALEOZOIC NONMARINE SNAILS

those found in Dendropupa vetusta (pi. 4, figs. 2-4).

The aperture of A. sturgeoni is remarkable primarily for the bulbous termination of the parietal barrier 

(pi. 9, figs. 1-3), the deep recession and size reduction 

in the columellar barrier (which is barely visible from 

the angle shown on pi. 9, fig. 1), and the recession of 

the palatal ridge (pi. 9, fig. 2). The weakly bifurcated 

shape of the basal ridge shows well from the low-angle 

view shown on plate 9, figure 3. A comparison of figures 3 and 4 of plate 9 with the growth sequences 

shown in figures 4 and 5 indicates that lip thickening 

and growth patterns in the A. sturgeoni and in A. 

ohioensis are similar.

Despite the fragmentary and limited material available, the differences in shell ribbing and barriers are 

substantial enough to warrant naming A. sturgeoni a 

distinct species. The basic patterns of lip formation, 

barrier position, and ridge formation suggest that only 

specific-level differentiation from A. ohioensis is involved.

We consider it appropriate to name this species 

after Myron T. Sturgeon, who has worked in the Dunkard beds and has contributed materially to knowledge of the late Paleozoic gastropods of Ohio.

Anthracopupa parva (Stauffer and Schroyer, 1920)

Plate 1, figures 15-17, 20-22

LoxonemaC!) parva Stauffer and Schroyer, 1920, p. 145, pi. 11, 

figs. 17-21.

We have examined the type specimens in the OSU 

Orton Museum, but because they lack some significant characteristics, we are uncertain as to the affinities of this species. All the specimens are juvenile, and 

only in the specimen we designate as the lectotype (pi. 

1, figs. 15, 20), can any part of the aperture be examined. We can detect no trace of any barriers on the 

parietal wall, and the columellar wall is partly broken 

and obscured, but has a trace of a barrier.

The sculpture of these specimens is quite similar to 

that of Anthracopupa sturgeoni (pi. 9, figs. 1-5) but is 

slightly finer. A. sturgeoni and ? A. parva differ most 

obviously in the spire angle; A. sturgeoni has a narrower apical angle and a slightly wider body whorl. In 

contrast, the spire angle of all specimens of ?A. parva 

is greater, and the lower whorls are distinctly wider 

than in A. sturgeoni. Until we have adult specimens of 

?A. parva, or at least juveniles having clear apertures 

so that the presence or absence of barriers can be 

checked, any generic assignment must be tentative. 

The spire angle of ? A. parva is intermediate between 

the wider angle of A. ohioensis and the very narrow 

angle of A. sturgeoni. The differences in these features are matched by the range of variation among

species of extant tornatellinid genera. Tentative inclusion of Loxonema (?) parva in Anthracopupa is a 

conservative and appropriate treatment.

Anthracopupa sturgeoni is known only from the 

Little Captina Limestone Member of the Monongahala Formation and "Uniontown Limestone Member" 

of the Uniontown Formation, whereas ?A. parva is 

from the "Lower Washington Limestone Member" of 

the Washington Formation. No specimens referrable 

to ?A. parva were taken by Yochelson in his fieldwork 

on the Dunkard and Monongahela Groups.

Anthracopupa vermilionensis (Bradley, 1872) 

Plate 1, figures 5-7

Pupa vermilionensis Bradley, 1872, p. 87, 88, fig. 1; Dawson, 1880, 

p. 410, 411, figs. 8, 9, 14c; White, 1883, p. 456, pi. 2, figs. 13, 14; 

Knight, 1941, p. 190, pi. 96, figs. 4a, b; Peck and McFarland, 

1954, p. 297, 306, pi. 29, fig. 4.

Maturipupa vermilionensis (Bradley), Pilsbry, 1926, p. 317, pi. 32, 

figs. 17,18; Henderson, 1935, p. 150,151; Yen, 1949, p. 236; Zilch, 

1959-1960, p. 64, fig. 203; Knight, Batten and Yochelson, 1960, 

p. 1318, fig. 211; Yochelson and Saunders, 1967, p. 125.

The location of the type specimens is not known to 

us. Scattered material in collections dating from the 

late 1800's and early 1900's probably consist of specimens collected by Bradley, if not indeed actually representing cotype material. Included in this array are 

a miscellany of specimens FMNH 51464 from the 

Walker Museum, FMNH 11675 from the Gurley collection, FMNH 4028 from the Illinois State Museum, 

and USNM 27611 received from F. H. Bradley in 

1898. Size variation in the seven adults^rom the above 

sets is: height of shell, 3.09-3.72 mm (mean 

3.35 ±0.087 mm), diameter, 1.55-1.84 mm (mean 

1.72 ±0.046 mm), height/diameter ratio 1.84-2.14 

(mean 1.94 ±0.042), whorls 4 7/s to 5% (mean 

5.08 ±0.114). The shells are thus slightly longer, are a 

little more slender, and have a very slightly higher 

whorl count than most of the measured Anthracopupa ohioensis (table 2). Although we maintain A. 

vermilionensis as a distinct species, collection of wellpreserved and more comprehensive materials might 

show that the populations fall within the range of A. 

ohioensis.

None of the specimens available to us had the palatal lip preserved, and most of the specimens had the 

aperture blocked by debris. Two partly cleaned specimens that had the palatal lip badly damaged or missing as a result of preparation showed the same parietal 

and columellar barriers as in A. ohioensis (pi. 1, fig. 5).

Some fragmentary specimens of A. vermilionensis 

are preserved in the Illinois State Geological Survey. 

Generally more specimens of Dawsonella than of A. 

vermilionensis are in these same samples, some of 

which contain many specimens of Spirorbis. A.
ANALYSIS AND CLASSIFICATION OF TAXA 25

ohioensis samples also contain many specimens of 

Spirorbis. None of the specimens loaned by Dr. Lois 

Kent had the apertures intact, and none add any new 

data to our understanding of the species. They do provide additional verified localities of occurrence: 

Freshwater beds below Macoupin, Center NV2, sec.

27, T. 7 N., R. 10 E., Jasper Co., Illinois. J. M.

Weller. 

St. David Formation,5 along Little Vermilion River, a

short distance above Petty's Ford, SE V4 NWV4, sec.

2, T. 17N., R.11W., Vermilion Co., Illinois. J. M.

Weller, 1928. No Spirorbis.

South bank of Hodges Creek, northwest of Chesterfield, SWV4 sec. 30, T. 10 N., R. 9 W., Macoupin Co.,

Illinois. J. M. Weller, 1927. 

West of coal mine in bluff on right bank, SWVi, sec.

29, T. 17 N., R. 10 W., Vermilion Co., Indiana. 

No count was attempted because of the multiple nature of the fragments and the thickness of the matrix. 

These records do confirm the joint occurrence of A. 

uermilionensis and Dawsonella, indicate that at least 

rarely they were found with Spirorbis, and provide 

potential sites for additional collecting activities.

Because of its shape, sculpture, and basic barrier 

structure, A. uermilionensis unquestionably belongs 

to the same genus as A. ohioensis, and it may well be 

conspecific with that species.

Subfamily DAWSONELLINAE Wenz, 1938

Description.—Typical neritoid shelf concave over 

umbilicus, spire normally elevated, few whorls without marked sculpture, lip weakly expanded and thickened.

Genus Dawsonella Bradley, 1874

Dawsonella Bradley, 1874, p. 151. Type species: Anomphalus

meeki Bradley, 1872; original designation.

Description.—Shell small, with as many as 3% 

rather rapidly widening whorls. Lip thickened and expanded, sharply angled at basal-columellar margin. 

Umbilicus covered in juveniles and adults by a concave callus that narrows the aperture. No prominent 

shell sculpture.

Discussion.—Dawson (1880, p. 413) compared 

Dawsonella with two polygyrid land snails, Stenotrema leai (Binney, 1841) and Triodopsis (Xolotrema) notata (Deshayes, 1830), but the barrierconstricted apertures of these shells, their strong surface sculpture, large number of whorls, reflected but 

thin adult lip, and large size (7-25 mm) easily distins "St. David Limestone Member" of the Carbondale Formation, as currently used in 

Illinois nomenclature.

guish them from Dawsonella and make any relationship to Dawsonella highly improbable. Whitfield 

(1881, p. 127, figs. 5,6) was the first to call attention to 

the similarities of Dawsonella to members of the family Helicinidae. The subsequent placement either in 

the Helicinidae or in a family unit, Dawsonellidae, 

standing next to the Helicinidae, is justifiable, but it is 

probably more conservative to treat the Paleozoic 

group as subordinate.

Dawsonella meeki (Bradley, 1872) 

Plate 10, figures 1-6

Anomphalus meeki Bradley, 1872, p. 88, fig. 2.

Dawsonella meeki (Bradley), Bradley, 1874, p. 151; Dawson, 1880, 

p.413, figs. 12, 13; Whitfield, 1881, p.127, figs. 5, 6; Henderson, 

1935, p. 159; Wenz, 1938, p. 52, 62, 434, 435, fig. 1070; Knight, 

1941, p. 97,98, pi. 96, figs, la-c; Yen, 1949, p. 236, 238; Peck and 

McFarland, 1954, p. 304, pi. 29, figs. 5,6; Knight, Batten, and 

Yochelson, 1960, p. 1279, fig. 189.3; Yochelson and Saunders, 

1967, p. 64; Solem, 1974, p. 175, fig. 3.

The location of the holotype is unknown. Six undistorted adult shells in FMNH 11673 from the Gurley 

Collection measured: height of shell, 2.30-2.89 mm 

(mean 2.66 mm), diameter, 3.78-4.11 mm (mean 3.99 

mm), height/diameter ratio, 0.565-0.710 (mean 

0.664), whorls, 3 1A to 3 5/s. They conform well to the 

material figured by Knight (1941) and Peck and 

McFarland (1954). Fragments in the Illinois State 

Geological Survey field collections made by J. M. 

Weller and listed under Anthracopupa uermilionensis 

were all subadult.

The most characteristic feature of the shell is a 

slightly concave plate (pi. 10, figs. 1, 2) that extends 

across the umbilical region. This plate is present in juveniles as well as adults, has an abrupt apertural margin, and effectively narrows the aperture. This plate is 

the typical neritacean shelf and, despite the lack of direct evidence, we assume it was accompanied by a neritoid operculum. Few dead helicinids that are found 

today in stream drift have the operculum in place, and 

indeed, few helicinid opercula have been found in 

drift deposits. The flat pebble shape of the operculum 

does not lend itself to accidental transport, whereas 

the globular shell can entrap a bubble of air and thus 

be transported for a considerable distance by a brief 

flood following a heavy rain. The absence of opercula 

in these deposits is not surprising to us.

The shell lip is slightly reflected (pi. 10, figs. 1,2), is 

only moderately thickened internally (pi. 10, fig. 1), 

and has a fairly heavy parietal callus (pi. 10, fig. 2). 

One highly significant feature relating to classification 

of this species is found at the basal-columellar margin 

(upper right of figs. 1, 2, of pi. 10) where the lip is 

sharply angled and protrudes in a right-angled turn.
26 NORTH AMERICAN PALEOZOIC LAND SNAILS, SUMMARY OF OTHER PALEOZOIC NONMARINE SNAILS

This protruded corner of the aperture is a characteristic of such modern helicinids as Pseudhelicina 

Sykes, 1907, Waldemaria A. J. Wagner, 1907, Oligyra 

Say, 1818, Alcadia Gray, 1840, Pleuropoma Moellendorff, 1893, and Schasicheila Shuttleworth, 1852. The 

heavy umbilical callus is a general feature of the Helicinidae, as is the low whorl count.

Surface sculpture of the shell (pi. 10, fig. 3) is undistinguished, showing no major features that would be 

indicative of a particular group. The shell structure itself (pi. 10, figs. 4-6) has a rather complex pattern of 

shell layers but the complex layers probably are a replacement phenomenon rather than the original 

structure.

Fragmentary specimens having shattered whorls 

show that the shell was quite thick and solid, another 

feature agreeing with the pattern seen in such drier 

zone helicinid species as Oligyra orbiculata tropica 

(Pfeiffer, 1852) and the Pacific Islands Orobophana 

Wagner, 1905, complex.

Because the shell of Dawsonella meeki is relatively 

simple in structure, it has few positive features to suggest family-level affinities. The umbilical callus, neritoid shelf, protruding basal-columellar lip junction, 

and thickening of the lip internally are all fully consistent with the extant Helicinidae. The basic shell 

shape, possession of only a few whorls, size, and lack of 

positive shell sculpture also are consistent with the 

Helicinidae. We can find no characters to separate 

Dawsonella from the Helicinidae.

Because of the concave umbilical callus surface and 

the large time gap, we accept the Dawsonellinae 

Wenz, 1938, as a subfamily unit of the Helicinidae.

OTHER PALEOZOIC NONMARINE SNAILS

We cannot offer a detailed review of Paleozoic nonmarine snails from other parts of the world. We do 

provide comments and suggestions for study, on the 

basis of a survey of the literature and examination of 

some materials. We try to place both the land and 

freshwater taxa within the systematic framework and 

the time scale worked out for the North American 

land taxa. Comments given by Yochelson (1975, p. 

254-256) concerning the placement of some of these 

species were preliminary and, in the main, were based 

on a misunderstanding of the genera.

A fair record exists of one species of land snail from 

England. Anthracopupa britannica Cox (1926, p. 

407-410, figs. 1,2) was described from the Keele beds 

(Westphalian D) near Hagley Park, Birmingham, 

England. The species was reassigned to Maturipupa 

by Yen (1949), but this reassignment has been ignored 

by later British workers. The outline drawings of the

original material show a well-developed parietal barrier and suggest a low, rather recessed columellar barrier equivalent to that described for Anthracopupa 

sturgeoni. In size and shape, A. brittanica is similar to 

A. ohioensis, but differs in possessing a lower, apparently deeply recessed, columellar barrier.

Poole (1969) summarized occurrences to date of the 

species in boreholes and indicated a local stratigraphic range of more than 350 feet (107 m). Calver 

(1969), in a definitive summary of the Westphalian of 

Britain, reported A. britannica from several localities 

in the Pennines and Midlands. At one locality, Uptown borehole, Oxfordshire, it ranges through about 

1,800 feet (549 m) of strata. Calver dated the beds 

then known to contain the species as probably late 

Westphalian D, but he could not rule out an early Stephanian age. Subsequently, both the geographic and 

geologic ranges have been extended significantly. 

Calver (in Richardson and Francis, 1971, p. 238, pi. 22, 

figs. 1-4) reported finding this species at half a dozen 

localities in the north of England and in southern 

Scotland, where he identified it in beds of Westphalian A age. His data included a few measurements and 

photographs of the species.

The single French record of a Paleozoic land snail is 

the specimen named Dendropupa walchiarum Fischer (1885, p. 100) from the "Permien moyen de 

Chambois (Saone-et-Loire)." This report describes a 

partial individual, estimated to have been 12 mm high 

and 4 mm in diameter when complete. All published 

information on shape, size, sculpture, and aperture 

are consistent with the structures reported above for 

Dendropupa vetusta. We have not been able to locate 

the holotype. It is not in the collection of the Journal 

de Conchyliologie, the Museum National d'Histoire 

Naturel, Paris, which holds many of the types of species described in that publication, nor in the Ecole de 

Mines. Without restudy of this unique fossil, we cannot comment further on its possible affinities. Pruvost 

(1957, p. 19) notes that Saone-et-Loire is in the Autun 

Basin. A geologic map (Feys and Greber, 1964, p. 49) 

shows Chambois to be near the north edge of the basin, north-northwest of Autun. It is in an area mapped 

as lower Autunian, in the Lower Permian. The record 

is thus much younger than that of Dendropupa vetusta, which is known from the lower upper Carboniferous. Whether this species is specifically or 

generically separable from Dendropupa is unknown.

The German Anthracopupa rothenburgensis 

Hintze (1933) from the Mansfelder Schichten of 

Halle, West Germany, is only slightly better known; 

the material is late late Carboniferous (early Stephanian) in age (H. Jaeger, written commun., 1971). As indicated by the published sketches, this is a slender,
OTHER PALEOZOIC NONMARINE SNAILS 27

rather small species that has three barriers in the aperture. The published data do not indicate whether 

the lip was thickened, and A. rothenburgensis may 

well prove to belong to a distinct genus. The age of A. 

rothenburgensis fits in well with the record of other 

species of Anthracopupa; A. rothenburgensis is intermediate between the age of A. ohioensis and that of A. 

vermilionensis. A small helicoid snail from the same 

deposit was not named. It is only 2.2 mm in maximum 

dimension and has a deep and widely open umbilicus. 

We can offer no suggestions as to its affinities.

Staesche (1963, 1969) discussed the fauna of the 

East German Lower Permian Rotliegende. Several 

forms had been called gastropods, but we agree with 

Dawson (1880) and Staesche (1963,1969) that none of 

this material can be considered of gastropod origin. 

Probably all are worm tubes, closely related to or 

identical with Spirorbis.

A potentially diverse fauna of land snails has been 

collected from the Lower Permian beds near Karniowice, 35 km west of Krakow, Poland. Dendropupa 

zarecznyi Panow (1936, p. 37-41, pi. 1, figs. 1-6) consists of two species. The larger (Panow, 1936, pi. 1, 

figs. 1,2) is a Dendropupa related to D. vetusta. Additional specimens from the type locality were collected 

in 1968 by E. L. Yochelson, S. Gasioroski, and A. 

Gromczakiewicz. Some years ago, specimens from 

Panow's collection were donated to the USNM. One 

steinkern in the USNM collection shows the same 

open umbilical region found in D. vetusta. On another 

specimen, fragments of axial sculpture are preserved. 

We have no doubt that these specimens are Dendropupa. Obvious differences from D. vetusta include a 

lower whorl count (less than seven), apparently wider 

whorls, and finer sculpture. We can offer no opinion as 

to the relationships among Dendropupa vetusta, D. 

walchiarum, and D. zarecznyi.

The smaller, narrower species (Panow, 1936, pi. 1, 

figs. 3-6) is less abundantly represented in the material available to us, and the specimens are slightly to 

moderately distorted. No traces of any sculpture remain, and we can only comment that in shape and 

form the specimens resemble Dendropupa primaeva 

(Matthew, 1895).

Yochelson collected his material in the lower part of 

the Karnovice Limestone, which is about 6m in maximum thickness and is exposed near a hillcrest. The 

few specimens were associated with large fragments of 

stems and leaves. Examination of the outcrop showed 

that the lower part of this unit contains abundant 

broken plants and leaves. The upper part has two lithologies that are repeated several times. One of these is 

an "organ pipe" limestone, apparently formed by lime 

deposited around the stems of reedlike plants, which

are now elongate holes in the rock. Alternating with 

this is ^ fine-grained limestone, locally having some 

holes, jvhich is suggestive of deposition by an algal 

mat. Although the Karnovice beds are conventionally 

referred to as a travertine, they might equally well be 

interpreted as a lime-rich shallow pond.

The late Dr. Jon Weber, of Pennsylvania State University,! kindly analyzed the isotopic composition of all 

three limestones. Data are reported in relation to the 

PDB standard in standard a, notation as:

C13 O18

loWer limestone _ _ _ 4.94 2.14 

"reed bed" ______ -6.54 -3.50

"alplbed" ______ -4.30 -1.81

i

He reported that these values are typical of freshwater 

carbonate rocks. With other evidence, they could be 

used to (support a conclusion in favor of freshwater depositioii for the Karnovice Limestone.

The kge of this deposit has been redefined as early 

Autunian by Lipiarski (1971) in her revision of the 

Carboniferous and Permian plants of Poland. Lipiarski uses Autunian as nearly equivalent to the German Rotliegende, the uppermost part of the unit 

being S|axonian.

During 1975, Yochelson was shown two European 

Lower Ipermian assemblages of land snails that have 

not yet been reported in the literature. Near Cesky 

Brod, east of Prague, Czechoslovakia, several thin 

limestones in a red-shale sequence contain specimens 

of an Anthracopupa and possibly of other taxa. In the 

vicinity | of Eggenberg, Austria, Anthracopupa occurs 

sparingly in similar rocks. These records will extend 

the geographic distribution of Anthracopupa, but we 

cannot <j)omment on specific affinities at this time.

Records of Paleozoic nonmarine snails from the 

U.S.S.Rj. are equivocal. Cox (1953) directed attention 

to the rfeport of Revunova (1938) on Tartarian Age 

snails fik>m the Sukhona-Mezen Rivers area, in the 

Vologdi region north-northwest of Moscow. They 

were described as freshwater taxa, but the late Dr. Boris Lich^irev (written commun., 1971) reported that 

"It is nejcessary for me to tell you that the species in 

my possession was described by Revunova in 1938 as 

Omphalpptychia malachovi. The author of that article did not clean the aperture of his specimens and 

missed tlhe presence of a tooth therein." Thus, at least 

some of this material may represent land-snail species, but without restudy, no additional comments can 

be made.

These are the only certain records of Old World Paleozoic land snails or possible land-snail fossils that we 

have bee|n able to locate. They extend the geographic 

ranges of Anthracopupa and Dendropupa from
28 NORTH AMERICAN PALEOZOIC LAND SNAILS, SUMMARY OF OTHER PALEOZOIC NONMARINE SNAILS

North America to Europe and also increase the known 

time range of Dendropupa into the Early Permian.

Several records of Paleozoic freshwater gastropods 

exist and are summarized below, although we can offer no meaningful comments on the affinities of the 

taxa. They do establish that the fresh waters had been 

colonized successfully by a variety of taxa in the late 

Paleozoic.

The earliest record is from Devonian beds near 

D'Aiguillon, Gaspe Peninsula, Quebec, Canada (Russell, 1947, p. 4). "A small gastropod, of generalized, viviparid-like form, was obtained" together with a clam 

similar to Modiomorpha, eurypterids, and fish remains. To our knowledge, this gastropod has not been 

described.

The species Carbonispira scotica Yen (1949) is represented by a single flattened, partly destroyed 

specimen. Although the specimen is dubiously describable and in poor condition, the suggestion by Yen 

that this may be a thiarid prosobranch is the best that 

can be made at present. The single specimen of Carbonispira was collected from shales below the Top 

Hosie Limestone of Namurian A, early late Carboniferous age (George and Black, 1971, p. 199).

Bernicia praecursor Cox (1927) was compared to 

the amnicolid hybrobiids, a conclusion with which we 

do not disagree. It is from rocks of the Scremerston 

Coal Group, of late Early Carboniferous (Visean D2) 

age (George and Black, 1971, p. 335), from near Scremerston, Northumberland, England.

Hydrobia gondwanica Cox (1953) from the Upper 

Permian Karroo Beds of Southern Rhodesia is an 

elongated hydrobiid shell and shows few unusual shell 

features. The existence of the European modern genus Hydrobia in the Permian is improbable, but proposal of a new generic name is unwise without added 

information on the shell structure. Subsequently, 

Rossouw (1970) reported, but neither described nor illustrated, a large (34 mm) freshwater snail from the 

Lower Beaufort beds at Weltevrede, South Africa, 

and an obscure early record of a planorbiform snail in 

the upper Paleozoic of Madagascar. The latter could 

be based on worm tubes.

Runnegar and Newell (1971, p. 25, 26, 54, figs. 24AC) have provided one record from the Lower Permian 

in the Estrada Nova Formation in the Parana basin of 

Brazil. Yochelson was quoted as referring these to 

Dendropupa, an error in judgment made at a time 

when Dendropupa was poorly understood. Restudy of 

these casts, from USGS locality 22836-PC, suggests 

that the material represents at least two, if not more, 

species of freshwater "hydrobiid" snails. The elongated morph that was figured by Runnegar and Newell (1971, figs. 24A-C) has an angled periphery, and, in

the better preserved large individuals, a truncated columella. This pattern is exactly equivalent to the modern hydrobiid Lyrodes Doering, 1884, in its 

nonspinate stage, and to the Old World genus Potamopyrgus Stimpson, 1865. The apparent columellar 

barrier mentioned by Runnegar and Newell (1971, p. 

26, fig. 24A) appears to be based on a fleck of dirt that 

was accentuated in retouching. The short, broader 

morph is roughly equivalent to such modern Neotropical hydrobiids as Aroapyrgus alleei Morrison (1946, 

pi. 2, fig. 4). In his review of the Tertiary nonmarine 

mollusks of South America, Parodiz (1969, p. 117,118) 

reported Lyrodes from the Oligocene of Colombia. 

Other taxa are Miocene and Pliocene, so that early 

Tertiary and Mesozoic records are lacking.

The fragmentary record of late Paleozoic freshwater snails includes apparent hydrobiids from England, Rhodesia, and Brazil; a possible thiarid from 

England; a "viviparid-like form" from Canada; and 

unclassifiable materials from South Africa and Madagascar. No freshwater pulmonate taxa have been recorded, despite the extensive freshwater bivalve fauna 

from many areas of the world. The data are insufficient to permit any sweeping conclusions. We can say 

that the record to date shows that freshwater gastropods preceded terrestrial ones by some millions of 

years. Paleozoic freshwater taxa occur both in Europe 

and in areas that formerly constituted Gondwanaland 

and they are found in older beds in the Northern 

Hemisphere. Terrestrial forms are known only from 

the Northern Hemisphere.

SYSTEMATIC SUMMARY

The classification of the Paleozoic land snails 

known from North America which has been discussed 

in the preceding pages is summarized in table 6; the 

ordinal and superfamily groupings are from the system proposed by Solem (1978), which in turn is modified from the systems of H. A. Pilsbry and H. B. 

Baker.

The distribution of the 10 fossil species among 5 

families and 3 orders of land snails emphasizes that a 

diverse fauna of land snails already existed before the 

end of the Paleozoic.

The confidence levels of family-group assignments 

for the studied material are summarized below. Solem 

judges that the evidence for family-level placement of 

Anthracopupa and Protodiscus is more than substantial. The complexity of the shell microstructural-ornament details in both fossil taxa and their extant 

relatives is such that convergent evolution is most improbable. We judge that these family assignments are 

quite firm.
PHYLETIC SIGNIFICANCE 29

TABLE 6. Systematic list of the Paleozoic land snails 

Class Gastropoda 

Subclass Prosobranchia 

Order Diotocardia

Superfamily Neritacea 

Family Helicinidae 

Subfamily Dawsonellinae

Dawsonella meeki (Bradley,

1872)

Subclass Pulmonata 

Superorder Stylommatophora 

Order Orthurethra

Superfamily Achatinellacea 

Family Tornatellinidae 

Subfamily Anthracopupinae Wenz, 

1938

Anthracopupa ohioensis Whitfield, 1881

?Anthracopupa parua 

(Stauffer and Schroyer, 

1920) 

Anthracopupa sturgeoni Solem and Yochelson, n.sp. 

Anthracopupa vermilionensis

(Bradley, 1872) 

Superfamily Pupillacea

"Pupa" bigsbii Dawson, 1880 

Superfamily Partulacea 

Family Enidae

Subfamily Dendropupinae Wenz, 

1938 

Dendropupa uetusta (Dawson,

1867) 

?Dendropupa grandaeva

(Dawson, 1880) 

Dendropupa primaeua

(Matthew, 1895) 

Order Sigmurethra 

Suborder Aulacopoda 

Superfamily Arionacea 

Family Discidae

Protodiscus priscus (Carpenter, 1867)

The evidence concerning Dawsonella and Dendropupa is at a somewhat lower confidence level. The 

shell structure of Dawsonella is relatively simple. All 

of its few unusual features agree with the structures 

found in the extant Helincinidae. These structures are 

not unique among extant gastropods, although the 

particular combination of structures seen in Dawsonella is not duplicated in any other family. We cannot 

rule out the possibility that Dawsonella is convergent 

to the Helicinidae, but because no structures of Dawsonella are incompatible with its assignment to the

helicinids, we are convinced that this family assignment is reasonable. Dendropupa shows most similarity to the Enidae and has been placed in that family. 

The genus could be a prototype for the Urocoptidae, 

but So|lem considers that this is unlikely, as basic 

shell-growth features in Dendropupa and the Urocop^ 

tidae ai^e inconsistant. Dendropupa may belong to an 

extinct1 family unit that was ancestral to the enid lineage, bjit the preponderance of evidence supports our 

including it as a subfamily of the Enidae.

The assignment of "Pupa" bigsbii to the Pupillacea 

is based on only one character complex, the apex and 

spire growth pattern. This pattern effectively separates this species from the Tornatellinidae, but assignment to any family unit of the Pupillacea is 

precarious. 

i

I PHYLETIC SIGNIFICANCE

The occurrence of five families- and three orders of 

land snails in the Pennsylvanian-Permian section of 

the fossil record has several interesting evolutionary 

implications. First, considerable taxonomic diversity 

is exhibited by the oldest known land snails. Indeed, 

half th^ gastropod orders that have produced land 

snails wfere present when this group initially appeared 

in the fiossil record. Second, all the five family-level 

groups found in the Paleozoic are still extant. This 

fact indicates a basic conservatism among the higher 

land-sn^il taxa. Third, the time and sequence of appearancfe of the gastropod orders have considerable 

bearing | on evolutionary direction and speculations as 

to interbrdinal relationships.

The qldest known Archaeogastropoda appeared in 

the Latb Cambrian. Yochelson (unpub. data, 1977) 

judges that all the assignments of Early and Middle 

Cambrian fossils to the Class Gastropoda that are recorded in the literature are spurious. The Superfamily 

Neritacea is recorded first from the Middle Devonian 

of Great Britian, about 375 million years ago. By the 

Early Mississippian, neritaceans were widely spread 

in marine deposits. Thus, presence of a terrestrial neritacean, Dawsonella, by the late Middle Pennsylvanian medicates that less than 75,000,000 years passed 

between! tne origin of the neritaceans and their recorded presence on land. Discovery of terrestrial neritaceans ! in earlier deposits would not be surprising. 

The earliest record of a freshwater neritacean, however, is nolf until the very end of the Jurassic (Mesoneritina Ye;n, 1946, from the Morrison Formation of 

western North America).

The presence of two orders and four families of the 

superorder Stylommatophora in the late Paleozoic is 

somewhat surprising. Conventionally, the superorder 

Basomniatophora has been assumed to be more primi-
30 NORTH AMERICAN PALEOZOIC LAND SNAILS, SUMMARY OF OTHER PALEOZOIC NONMARINE SNAILS

tive than and ancestral to the superorder Stylommatophora. However, the first terrestrial basommatophoran is the ellobiid genus Carychium Miiller, 

1774, which appeared in the Late Jurassic of Europe; 

the first freshwater ellobiids are from the uppermost 

part of the Upper Jurassic Morrison Formation (Tortacella White, 1895, Zaptychius Walcott, 1883, Mesochilina Yen, 1951, Mesauriculstra Yen, 1952); the 

first marine ellobiids seem to be from the Upper Jurassic of Europe; and the first appearance of the higher limnic Basommatophora is in the same Morrison 

Formation (Lymnaeidae, Planorbidae, Physidae).

Thus, the superorder Basommatophora appeared 

about 150 million years later than the Stylommatophora. Perhaps the Basommatophora was derived 

from the Stylommatophora, the opposite of current 

thinking, or perhaps each had an independent origin 

from more primitive stocks.

In our opinion, the reality of this time gap cannot be 

seriously questioned. Even though the time of first appearance in the fossil record and the time of origin are 

not the same, the first records of both superorders indicate that equivalent degrees of diversity had been 

reached. Thus, pushing back the origin of the Basommatophora, could entail a similar backwards shift of 

the origin of the Stylommatophora. If the freshwater 

mollusk record were essentially unknown before the 

Late Jurassic, the absence of Basommatophora might 

not be considered highly significant. However, a scattered, but significant, record of early freshwater mollusks does exist. The Paleozoic viviparid-form, 

hydrobiid-form, and thiarid-like records were noted 

above. The Valvatidae and the Viviparidae are known 

from the Triassic, along with a very extensive record 

of freshwater clams. The earliest non-marine bivalve 

group seems to be the Late Devonian family Archanodontidae Weir, 1969, from the United Kingdom and 

North America, based on Archanodon Howse, 1878 

(-Amnigenia Hall, 1885). By the early Carboniferous, 

a diverse group of "unionid precursors" existed, even 

though the Unionidae themselves did not appear until 

the Triassic. The late Carboniferous also saw the family Myalinidae make an incursion into fresh waters 

(data supplied by John Pojeta, Jr., 1975).

Thus, a significant record exists of several freshwater taxa older than the Late Jurassic. The number 

of records is such that, very probably, basommatophorans would have been preserved if they had been 

present.

The Morrison Formation (Yen and Reeside, 1946; 

Yen, 1952) is thus viewed as an indicator of the first 

flourish of the Basommatophora and the first extensive documentation of a rich freshwater mollusk. Besides the1 Physidae, Lymnaeidae, and Planorbidae, the 

Morrison contains representatives of the Neritinidae 

(Mesoneritina Yen, 1946), "Hydrobiidae," Viviparidae, Valvatidae (Amplovalvata Yen, 1952), several 

"Unio, s. L," Vetulonaia Branson, 1935, and the enigmatic clam genus Hadrodon Yen, 1952. Limopsis Yen, 

1952, was referred to the Otinidae, but we suggest that 

evidence is insufficient to refer this taxon to a family 

unit. The only unusual added element in this assemblage was the large number of freshwater Ellobiidae, 

because this taxon today inhabits mangrove and nipa 

palm swamps on the coast or brackish-water mudflat 

areas, and does not live in freshwater. Apparently the 

Ellobiidae underwent an extensive late Mesozoic 

freshwater radiation.

The presence of freshwater clams and prosobranches in Paleozoic deposits shows that the fresh 

water had been colonized by mollusks and that conditions favorable for their preservation existed long before the first recorded appearance of the 

Basommatophora. Thus, the absence of basommatophorans from the fossil record until the Jurassic-Cretaceous boundary suggests that they evolved later 

rather than that chance prevented their preservation. 

The earlier appearance, and thus probable much earlier evolution, of the superorder Stylommatophora indicates the need to reevaluate the relationship 

between these two superorders and the previously hypothesized direction of evolution leading from the Basommatophora to the Stylommatophora.

We cannot resist speculating on one highly intriguing evolutionary correlation. The first appearance of 

the typical freshwater basommatophoran snails was at 

the end of the Jurassic in the Morrison Formation. 

Present in this fauna are the typical taxa of temporary 

ponds and streams, the families Physidae, Lymnaeidae, and Planorbidae. All these are capable of self-fertilization to start a new colony and frequently have 

been found on the feet and feathers of waterfowl; they 

are thus known to be dispersed by birds.

The coincidence of the sudden appearance of these 

freshwater snail taxa with the origin and radiation of 

birds cannot be ignored. As summarized by Cracraft 

(1973, p. 495, 496), the fabled Archaeopteryx appeared in the Late Jurassic, Australian and European 

records of waterfowl in the Early Cretaceous exist, 

and by the Late Cretaceous, a worldwide fauna of diverse waterfowl had evolved. Cracraft (1973, p. 496) 

concluded that "*** the Class Aves, much as we know 

it today, was present on all continents in the Cretaceous***" and that by the end of the Eocene ^^approximately 26 of the 32 known orders are found as 

fossils." Thus, the sudden appearance of the hitchhiking basommatophoran pond snails may have been the
ECOLOGICAL SIGNIFICANCE 31

result of pond-hopping birds in the Late Jurassic.

The essentially simultaneous recording of four families and two orders of the Stylommatophora sheds no 

light on the problem of origin and early evolution of 

the group. The fact that the fossil genera are referable 

to extant families with very high confidence limits 

(Anthracopupa and Protodiscus) or high confidence 

limits (Dawsonella and Dendropupa) indicates that 

by the first appearance in the fossil record of the Stylommatophora, their radiation already had reached a 

plane of stability. The Tornatellinidae (Anthracopupa), although complex in many features, have a 

number of generalized anatomical features and are 

placed as perhaps the most generalized group within 

the Orthuretha. In contrast, the Enidae (Dendropupa) show many advanced features, and some taxa 

have evolved "pseudosigmurethrous" features, which 

apparently have enabled them to colonize semiarid 

areas successfully. The Enidae are thus probably one 

of the most advanced families of the Orthurethra. The 

superfamily Pupillacea ("Pupa" bigsbii) is yet a third 

family-level group of the Orthurethra.

The fourth taxon of the Stylommatophora found in 

the late Paleozoic, the family Discidae, belongs to the 

arionid complex, which is one of the most primitive 

groups of the Order Sigmurethra. The Discidae is 

probably one of the more advanced families of the 

Arionacea, but it still belongs to a probable stem 

group of the Sigmurethra. The above data on landsnail phylogeny are summarized from Solem (1978b).

Possible reasons for the absence of the remaining 

land-snail ordinal groupings are worth considering. 

The absence of both freshwater and terrestrial basommatophoran snails until the Jurassic-Cretaceous 

boundary, despite the recording of freshwater mollusks since the Devonian, suggests a later period of 

evolution. The land-snail order Mesurethra, the suborder Holopodopes, and the suborder Holopoda (the 

latter two are subdivisions of the Sigmurethra) all 

first appear in the Upper Cretaceous of Europe. These 

fossils, like the Paleozoic taxa, also are assignable to 

extant families, and they indicate that a high level of 

differentiation had been achieved and stability 

reached by the time of their appearance. In general, 

most of these taxa have efficient water-conservation 

devices; thus, they are able to inhabit drier areas than 

the members of the order Orthurethra (except the 

arid-zone pupillaceans and enids) and the superfamily 

Arionacea. Most arionids still inhabit flood plains or 

very moist forests. They, and the terrestrial prosobranchs such as the Helicinidae, can be active only 

during periods of nearly saturated humidity. In contrast, the higher Sigmurethra, because they evolved

better water-conservation mechanisms, can be active 

in drier conditions and can more effectively retreat 

from both floods and drying niches. Hence the higher 

Sigmurethra would be less likely to be preserved as 

fossils than would the flood-plain-inhabiting Orthurethra and Arionacea. Exceptions to the above generalities occur, but differences in both habitat selection 

and mobility result in different chances of successful 

preservation for the ordinal groups that are present 

and for those that are absent from the Paleozoic deposits. The absence to date of land-snail fossils from 

the Early Permian to the Late Jurassic might well 

conceal the actual evolution of the missing ordinal 

units; we cannot exclude the possibility that higher 

forms were present in the late Paleozoic but were not 

preserved in the limited record because of habitat selection.

ECOLOGICAL SIGNIFICANCE

We can offer a few observations and speculations on 

the probable ecology of the Paleozoic land snails. The 

most abundant materials are those of Anthracopupa 

in the Little Captina Limestone Member of the Monongahela Formation and Dendropupa in the tree 

stumps of the Joggins Formation. Dawsonella is 

abundant in restricted Illinois localities. All other 

specimens are rare in comparison.

The occurrence of fresh and worn fossils of Anthracopupa in the same locality finds modern equivalents 

among pupillid land snails of North America. Streamdrift windrows in late spring often contain material of 

the same species in two conditions. Specimens without periostracum whose surfaces have been worn or 

eroded are mingled with fresh undamaged specimens. 

The former, after death, may have sat in standing water with decaying leaves for a short to medium period. 

The weak humic acids and (or) bacterial action could 

have removed the periostracal covering. When a subsequent rain storm washed them into a stream, they 

could have been joined by individuals that died at the 

same time but then laid in dry leaf litter and did not 

lose the periostracum or by live specimens that, after 

being washed into the stream and stranded, died on 

the spot. The thanatocoenoses of Anthracopupa 

could have originated in a similar way.

The postulated method of deposition is fully consistent with conclusions reached through study of the 

lithology and stratigraphy of the Dunkard beds (Berryhill, 1963, p. 80-90). Seasonally high rainfall could 

have flushed downstream the dead snails accumulated during preceding drier periods. A bubble of air inside the shell, trapped and held by the protruding 

apertural barriers, could have enabled the shell to
32 NORTH AMERICAN PALEOZOIC LAND SNAILS, SUMMARY OF OTHER PALEOZOIC NONMARINE SNAILS

float for a long time. Only breaking of the shell to release the air bubble or gradual filtering in of fine sediment would have reduced buoyancy and caused the 

shell to sink to the bottom. Wind drift, or eddy effects 

along stream margins could have caused the tiny 

shells to accumulate in shore drift and thus be subject 

to burial and preservation during the next rainstorms. 

This theory accounts for the presence of minute land 

snails in such drift deposits. The observed paucity or 

absence of larger shells in the drift deposits is also 

consistent with this environment. A larger shell, 6 mm 

or more in diameter, or a shell without any apertural 

barriers is far more apt to have any air bubble escape 

and thus lose buoyancy. In addition, the larger shells 

can more easily crash into one another and break, 

whereas the surface tension of the water film tends 

to keep the minute specimens apart and thus undamaged. This inference is supported by the high 

percentage of the large snails represented in drift deposits by broken fragments.

Carroll (1967, p. 112) has postulated that the standing fern trees of the Joggins were inundated by heavy 

floods and that vertebrates wandered onto the drying 

mud and into the decaying, partly buried tree trunks. 

A second flood would then bury and preserve the unlucky colonizers. Massive deposits of fragments of 

Dendropupa are consistent with this scenario, particularly if Dendropupa, rather than being a leaf-litter inhabitant, might have been associated with the 

tree-trunk-base debris in the fern forests. Such an 

ecological niche would be consistent with that of modern terrestrial snail taxa of equivalent size.

A possible present-day analog to the Dendropupa 

thanatocoenose is present in modern tropical forests. 

There, a variety of cyclindrical land snails 8-20 mm 

long inhabit the buttress and tangled root litter accumulations of large trees. The environment is somewhat acidic, turnover rates are high, and dead shells 

fragment quickly. The soil generally is thin and gravelly, so that runoff after heavy rains tumbles dead 

shells and gravel fragments, further breaking the 

shells. Accumulation of these in rock crevices, streamside drift, low tree holes, or in a buttress of a tree are 

all common phenomena.

In general, the taxa that have depositional associations in freshwater limestones similar to that of 

Anthracopupa, live today in open loose litter, under 

bark of decaying logs, and in or slightly above floodplain levels. They do not live in swamps or in continuously inundated areas, but in the seasonally flooded 

areas and humid forests adjacent to the flood levels. 

Some taxa have adapted to drier plains habitats, but 

the time their adaptation occurred is unknown. The 

tree-associated larger shelled taxa of the tropics today

are found mostly in monsoon or rain forest areas. We 

cannot assume from the snails that the Joggins section 

had a higher and less seasonal rainfall regime than the 

Dunkard. However, the snails and repeated widespread thin limestones of the Dunkard may indicate 

that the limestones were deposited in an area having a 

mildly seasonal rainfall pattern.

Land snails of the modern helicinid genus Oligyra 

commonly compose a large proportion of the streamdrift deposits in Texas and drier parts of Mexico. Apparently they can be transported tens if not hundreds 

of kilometers during periodic floods resulting from 

cloud bursts, as the thick shells are less easily 

damaged than the more fragile shells of the pupillid 

taxa also found in the semiarid regions of Texas and 

Mexico. The shells of Oligyra are common in these deposits, but their opercula are rarely found. An air bubble caught in the shell permits long-distance 

transport, whereas the flat solid operculum is far less 

apt to be stream transported.

Thus, all three of the abundant genera of Paleozoic 

land snails can be matched with modern equivalents 

in terms of probable ecology and chances for preservation. All three were transported moderate to long distances by floods. Although most frequently such 

transportation today results in the transport of dead 

shells or in the death of any live snails picked up by 

the floodwaters, occasionally to frequently, floodwaters successfully transfer live snails from one area to a 

site many kilometers downstream. Establishment of 

new colonies by stream transport is one of the more 

usual ways in which flood-plain land snails are dispersed. Such transport can, of course, happen only in 

a downstream direction. Even local movement of land 

snails can lead, in a relatively short time, to great 

range extensions. The last major glaciation in North 

America extended as far south as central Indiana and 

Illinois as late as 25,000 years ago. Yet land-snail species such as Vitrina limpida (Gould), Discus cronkhitei (Newcomb), Punctum minutissionum (Lea), 

Vertigo ventricosa (Morse), and Cionella lubrica 

(Miiller) have managed to reach the shores of Hudson 

Bay, a distance of at least 2,000 km, since the retreat 

of the glacier. Such movement has been basically 

against streamflow, so that flooding transport has not 

been a positive factor in their dispersal.

Although we present no new botanical data, speculation as to the factors that permitted the initial explosive radiation of land snails is very much in order. 

The first appearance of a diversity of land snails in the 

Joggins section (late early Pennsylvanian) of Nova 

Scotia is highly dramatic and was a point emphasized 

repeatedly by Sir William Dawson. The limited paleobotanical data do not identify any dramatic shift in
BIOGEOGRAPHIC SIGNIFICANCE 33

vegetation types that coincides with or slightly precedes this gastropod crawl onto land, but three trends 

in the evolution of Euramerican flora at about this 

time seem to have provided, in combination, conditions favorable to the colonization of land by snails.

The three floristic trends were the increase in arborescent plants at the expense of herbaceous types, 

the development of laminate foliage types in several 

lineages, and (associated with the development of 

laminate foliage types) development of a continuous 

leaf-shedding pattern typical of modern evergreen 

plants in addition to the already existing seasonal 

leaf-shedding pattern typical of modern deciduous 

plants (T. Phillips, oral commun., 1976). Although the 

arborescent pattern of growth is known to have been 

present by the Late Devonian, it did not become common until the latter part of the Mississippian. Plants 

showing these trends gradually became more common 

through the Mississippian.

The combined net effect of these changes for potential litter dwellers would have been considerable. 

Broadening of the higher leaves provides shade and 

substantially retards the dessication caused by exposure to direct sunlight. Broad leaves also add, through 

transpiration, considerable moisture to the shaded air 

layers. The trend towards continuous leaf shedding 

has two effects: continual addition of raw material to 

the litter and continual provision of better shade 

cover for the litter. These relatively simple trends 

would have combined to provide a moist, well-shaded, 

and continuously replenished litter a favorable environment for moisture-loving snails. Thus, the sudden 

explosive appearance of land snails in the Westphalian B could have been triggered by the above-mentioned botanical trends that produced a suitable 

continuously moist litter habitat. In contrast, at the 

same time, the Gondwana and Cathaysean floras were 

still shedding leaves seasonally rather than continuously. Thus, the fact that the first land snails found in 

the fossil record of the Gondwana areas are much 

younger than those found in the Joggins section might 

indicate that the land there was not favorable to snail 

colonization until the flora changed. However, this 

idea pushes our speculations to the outer limit.

Careful attention to earlier limestones that may 

have been deposited in freshwater ponds, the inferred 

habitat of the Anthracopupa assemblages, would provide the best chance of locating land-snail fossils older 

than those in the Joggins section. The chances for successful preservation of snails in these habitats would 

be much greater than those for the very unusual preservation of snails in tree stumps found in the Joggins. 

We would not be surprised if study of such earlier 

limestones did reveal traces of earlier land snails, but

we are not optimistic that this hunt will prove successful. Breakdown of litter to a size easily attacked by 

bacteria and fungi existed on land at the death of the 

earliest herbaceous plants. However, dry seasons, the 

absence of moisture-preserving shelter sites, seasonal 

rather than continuous additions to the litter, and resulting moisture-deficit problems would effectively 

have prevented colonization by most snails. To what 

extent the first land plants were restricted to continuously moist habitats or adjusted to seasonality of dryness is unknown, as is the identity of the early plantdetritus feeders. The Visean may well have been the 

first suitable opportunity for snails to colonize the 

land, and the Westphalian B records may represent 

the initial surge of land-snail taxa.

BIOGEOGRAPHIC SIGNIFICANCE

The occurrence of Anthracopupa in an area from 

Vienna, Austria, to the Illinois-Indiana border and the 

apparent presence of Dendropupa in eastern Canada, 

Poland, and France provide additional evidence that 

Europe and North America were joined together during the late Paleozoic. During the approximately 

75,000,000 years separating the earliest and latest records for Anthracopupa, it could easily have dispersed across this range of nearly 5,500 km. The 

dispersal of land snails across 2,000 km in North 

America during less than 25,000 years is evidence 

enough of their dispersal powers. We also would point 

out that bird (or even pterodactyl) accidental transport was not available to the Pennsylvanian land 

snails. Although some freshwater prosobranch snails 

have been known to emerge alive from the crop of a 

duck, and thus could be transported in bird stomachs 

as well as on bird feet, no pulmonate snails, which lack 

the protective operculum, have been known to undertake a similarly safe passage. The comparatively rare 

giant insects of the Pennsylvanian coal forests conceivably could have accidentally transported water 

snails, but resorting to such helpful chauffeurs is unnecessary to explain the distributions.

The comparative distributions of the Paleozoic and 

appropriate modern taxa are highly diverse. We discuss these in phylogenetic order. The modern Helicinidae are basically tropical and markedly disjunct in 

distribution; Southeast Asia to Polynesia and Central 

America and the West Indies are the two centers of 

diversity. A few genera, such as Pseudhelicinia, are 

endemic to South America, and a few species live in 

Florida or the southeast United States (Helicina Lamarck, 1799, Oligyra Say, 1818, and Lucidella Swainson, 1840). One genus, Hendersonia A. J. Wagner, 

1907, is a Holarctic relic that still can be found in scat-
34 NORTH AMERICAN PALEOZOIC LAND SNAILS, SUMMARY OF OTHER PALEOZOIC NONMARINE SNAILS

tered colonies through northern Iowa, southern Minnesota, Wisconsin, Michigan, and Illinois. From the 

structure of the operculum and the radula, Hendersonia is considered to be a primitive helicinid, somewhat related to the Japanese endemic Waldemaria. 

Fossil species assigned to Hendersonia are known 

from the Paleocene of Wyoming and Miocene of Oregon (Pilsbry, 1948, p. 1087). Thus, the presence of 

Dawsonella in Illinois during the late Paleozoic falls 

within the fringes of current geographic range.

The Tornatellinidae, except those transported by 

humans, are currently limited to the islands of Polynesia, Micronesia, and Juan Fernandez (see Cooke 

and Kondo, 1960, p. 22-31). The centers of speciation 

are the Austral Islands (35 species), Hawaii (152 species), and Juan Fernandez (19 species). Only a few 

small arboreal genera and species are widely distributed. There is a "***tenuity of relationship between 

the larger groups that indicates either great age and 

long isolation of the groups or destruction of the direct 

links leading to the highly variegated forms of today, 

or both" (Cooke and Kondo, 1960, p. 45). The discovery of late Paleozoic Anthracopupa in Illinois, Ohio, 

Pennsylvania, Nova Scotia, and Europe is thus quite 

an extension in geographic range. The only other 

known fossil tornatellinid is Protornatellina isoclina 

(White, 1895) from the Cretaceous of Wyoming. Polynesia today is an important center of snail diversity, 

harboring the relic land-snail families Endodontidae, 

Partulidae, and Amastridae and the achatinellid-tornatellinid complex.

The Pupillidae and other vertiginid families are 

mostly worldwide in distribution. No meaningful observations of changed distribution are possible, particularly in view of the tentative nature of our 

classifying "Pupa" bigsbii in the Pupillacea and its 

unique occurrence.

The Enidae are distributed in the Palearctic, China, 

India, and marginally southeast Asia and Indonesia; 

single taxa are in northern Australia, New Hebrides, 

and New Caledonia. A modest radiation in Africa 

completes their modern known range. The Nova Scotia record presents quite a different picture. It raises 

questions as to which group might have displaced the 

Enidae from North America, what group is the modern ecological equivalent in North America, and what 

special niche is occupied by the enids in the Old 

World. We do not have satisfactory answers to these 

questions and can only point them out as worthy of 

investigation.

Today the Discidae is a strictly Holarctic group containing only a few species. The greatest generic diversity is in North America, where humid montane areas 

or at least hill country contain the greatest variety of

forms. Essentially no shift in distribution has taken 

place since Protodiscus evolved.

To summarize, the changes in biogeographic patterns from late Paleozoic to Holocene range over a 

spectrum; they include no change (Discidae); worldwide distribution (Pupillacea); slight southern shift 

and separation into two centers of diversity, one in the 

Eastern Hemisphere and one in the Western Hemisphere (Helicinidae); displacement from North America to the Old World (Enidae); and isolation today on 

the outer islands of the Pacific (Tornatellinidae). 

More detailed discussion of comparative distribution 

through time has been given by Solem (1977).

REGISTER OF LOCALITIES FOR 

MONONGAHELA-DUNKARD GASTROPODS

The occurrence of well-preserved gastropods at 

many of localities in the northern part of the Dunkard 

basin was reported by Yochelson (1975). His list by no 

means gives all the localities at which lands snails may 

be found but is representative of the geographic distribution of these fossils.

Repeated below are locality data for those specimens from U.S. Geological Survey collections that 

have been illustrated or measured. Localities for additional collections of land snails in the Field Museum 

of Natural History, Chicago, are given here.

U.S. Geological Survey

24827-PC Clark Hill section of Cross (1950). Poorly 

exposed limestone in roadside ditch at 

altitude of approximately 1,300 feet 

(396 m), just below crest of hill, center 

SW !/4 sec. 16, Salem Twp., Monroe 

County, Ohio. Nineveh Limestone 

Member, Green Formation, possibly 

unit 21.

24836-PC In ravine on north side of Deep Run 

Road, extreme west sec. 26, Pease 

Twp., Belmont County, Ohio. Basal 

part Little Captina Limestone Member, Monongahela Formation.

24840-PC Pasture on south side of Ohio Route 148, 

just east of junction with Ohio Route 

145, in gully below bench, 30 m south of 

road, center of west line, sec. 28, Washington Twp., Belmont County, Ohio. 

Little Captina Limestone Member, 

Monongahela Formation.

24846-PC Ayers limestone quarry in SWVi sec. 19, 

Pease Twp., Belmont County Ohio.
REGISTER OF LOCALITIES 35

Thin limestone 0.6 m below base of 

main bed of "Lower Washington Limestone Member," Washington Formation.

24850-PC Pasture on south side of Ohio Route 148, 

center of west line, sec. 28, Washington 

Twp., Belmont County, Ohio (re-collection of 24840-PC). Little Captina 

Limestone Member, Monongahela 

Formation.

24851-PC North side of cut on unnamed secondary 

road east of Henderson Avenue, (highway 18), 0.68 km southwest of sewagedisposal plant, Washington West 

quad., Washington County, Pa. Uniontown Formation, Monongahela Group.

24854-PC In the bank of Deep Run on northwest 

side of county road 2, 0.72 km east of 

Ohio Route 798, Belmont County, Ohio 

(re-collection of 24836-PC). Little 

Captina Limestone Member, Monongahela Formation.

24855-PC Ayers limestone quarry in SWVi sec. 19, 

Pease Twp., Belmont County, Ohio (recollection of 24846-PC). "Lower Washington Limestone Member," Washington Formation.

24856-PC In bank on south side U.S. Route 40 behind old house at "S" bridge which is 

also point of divergence of Pa. Route 

221, Washington County, Pa. "Lower 

Washington Limestone Member," 

Washington Formation.

24857-PC B&O Railroad cut at Washington, Pa., 1.1 

km northwest of mosque in Washington Cemetery, Washington County, Pa. 

"Lower Washington Limestone Member," Washington Formation.

Field Museum of Natural History

[Description of locality, collector, date of collection. 

Comments on possible geologic occurrence are given in brackets.]

1-1 Scott's Cut on the B&O Railroad) about 1 mile 

(1.6 km) northwest of Warnock, Belmont 

County, Ohio. Lowest limestone bed (on the 

east side of the cut). P.O. Moore. July 4-6, 

1925. [Upper part of Monongahela Group, 

possibly Uniontown Formation.]

1-2 Scott's Cut (on the B&O Railroad) about 1 mile 

(1.6 km) northwest of Warnock, Belmont 

County, Ohio. Second limestone bed containing fossils in this cut, about 25 feet (7.6 m), 

above the level of the tracks and beneath the

shale zone that underlies the coal. P. D. 

Moore. July 6, 1925.

1-2 Same locality as above, but "from the middle limestone beds." [upper part of Monongahela 

Group, possibly Uniontown Formation or 

Waynesburg Formation of Dunkard Group.]

1-3 About 1 mile (or 1.6 km) northwest of Warnock, 

Belmont County, Ohio. Scott's Cut (on the 

B&O Railroad). From the black shale beneath 

the Waynesburg coal. July 8, 1925. [Probably 

part of the Gilboy Sandstone Member, Monongahela Formation.]

1-4 Along north side of Cadiz Road in bank beside 

road, !/4 mile (0.4 km) northwest of the village 

of Colerain, in southwest cor. SE 1A sec. 14, 

Colerain Twp., Belmont County, Ohio. Altitude 1,300 feet (396 m). "Upper Washington 

limestone" (according to Wilbur Stout). Wilbur Stout, Geo. White, and P. D. Moore. July 

1925. [Upper Washington Limestone Member," Washington Formation.]

1-4 Same locality as 1-4 above. From "Lower Washington limestone." P. D. Moore. July 7,1925. 

[Probably "Lower Washington Limestone 

Member," Washington Formation.]

1-5 Near center sec. 30, T 1 N, R 9 W, about % mile 

(1.2 km) southwest of Constitution, Washington County, Ohio. At an altitude of about 985- 

990 ft. (300.2-301.8 m) in the very top of the 

amphitheatre facing to the northwest. From 

the outcrop of the "Nineveh limestone" at the 

above place. The limestone here is about 10 in. 

(25.4 cm) thick and crops out for about 200 yd. 

(183 m) around the head of this sharp valley. 

Prof. R. W. Whipple and P. D. Moore. July 10, 

1925 [Correlations in the upper part of the 

Greene Formation are uncertain; this may not 

be the Nineveh Limestone Member.]

1-6 From outcrop in bed of the small tributary, 

south of Cat Run and west of the old byroad in 

the west-central part of sec. 19, York Twp., 

Belmont County Ohio. From the black limy 

shale about 6 ft. 4 in. (1.9 m) above the "Fulton green shale" outcrop in this gully. P. D. 

Moore. July 8,1925. [Probably from the Little 

Captina Limestone Member, Monongahela 

Formation.]

1-7 The very top of the little hill just east of the center SEV4 SWV4 sec. 27, T 2 N, R 9 W; about 6 

miles (9.6 km) southwest of Marietta, Ohio. 

South of the old railroad and from just inside 

the fork of the roads about '/i mile (0.4 km) 

east of the above location, in the pasture; also 

along the road in the north-central part of
36 NORTH AMERICAN PALEOZOIC LAND SNAILS, SUMMARY OF OTHER PALEOZOIC NONMARINE SNAILS

sec. 21. Prof. R.W. Whipple and P. D. 

Moore. July 13, 1925 [Possibly Dunkard 

Group.]

1-8 From limestone exposure in road 2/5 way up hill, 

on east bank of Muskingum River, 1 Vz miles 

(2.4 km) southeast of the mouth of Bear 

Creek, northwest of Marietta, Ohio. Some 

specimens are from boulders in creek bed. 

This is most probably No. 9 of the section given by Stauffer and Schroyer (1920, p. 127) 

"Section of east bank of Muskingum River, 

1 Vz miles southwest of mouth of Bear Creek." 

P.D. Moore. July 12,1925 [Near the top of the 

Monongahela Group.]

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INDEX

[Italic page numbers indicate both major references and descriptions]

Page

Abida————————————————22

Acadian Geology—————————————————————— 6 

Achatinella —————————————————— 22, 23

Achatinellacea ——————————————————————— 29 

Achatinellid-tornatellinid complex——————————34 

Achatinidae ————————————————————— 12, 21 

Acknowledgments ————————————————————— 2 

Afriboysidia —————————————————— 22

Alcadia—————————————————————26

Algal mat——————————————————————————27 

alleei, Aroapyrgus ————————————————28

Amastridae ———————————————————————— 34 

Ammgenia———————————————————— 30

Amplovalvata —————————————————— 30

Anauchen ——————————————————————22

Anguispira ———————————————————— 9

picta———————————————— 8, 9; pi. 3

sculpture, apical ——————————————— 8, 9 

Anomphalus meeki ———————————————— 25

Anthracopupa ————— 2, 3, 5, 9, 10, 20, 24, 27, 28, 31 

affinities————————————————————— 19, 20 

assemblages, habitat————————————————33 

bigsbii———————————————9

britannica ——————————————— 23, 26

collections, Ohio ——————————————————— 1 

compared to Cyclophoridae————————————20 

compared to Dendropupa————————— 20, 21

compared to Ellobiidae———————————————21 

compared to Stylommatophora—————————21 

compared to Tornatellinidae———————————22 

depositional associations ————————————— 32 

dispersal————————————————————————33 

dunkardana ———————————3, 5_ 15; pi. i

fossils —————————————————————————31 

geographic range —————————————27, 28, 34 

Illinois-Indiana border ———————————————33 

juvenile————————————————————19, 22, 23 

mode of preservation ————————————————19 

ohioensis ————————— 3, 4, 5, 15, 20, 23, 24,

25, 26, 27, 29; pis. 1, 5, 

f), 7,9

measurements —————————————————16 

mode of preservation —————————————18 

shape variation—————————————— 16, 17 

shell structure ——————————————————16 

size variation ——————————————— 16, 17 

variability————————————————— 16, 17 

parua————————————— 23, 24, 29; pi. 1

rothenburgensis———————————— 23, 26, 27

sturgeoni ——————————— 2, 2,'i, 26, 29; pi. 9

thanatocoenoses ———————————————————31 

uermilionensis ——————— 23, 24, 25, 27, 29; pi. 1 

Vienna, Austria———————————————————33 

sp ——————————————————————————pi 8

Anthracopupinae ————————————————— 3, 20, 29 

Archaeogastropoda ———————————————————29

Archaeopteryx—————————————————— ;j()

Archaeozonites priscus——————————————— 7

Archanodon —————————————————— 30

Archanodontidae ————————————————————— 30 

Arionacea —————————————————————— 29, 31 

Aroapyrgus alleei —————————————————— 28

Aulacopoda—————————————————————————29 

Aulacospira————————————————————22

Auriculella————————————————————23

Page 

Auriculidae ———————————————————— 20

Austral Islands ——————————————————————34 

Ayers limestone quarry, Belmont

County, Ohio ——————————————18, 34, 35 

specimens ———————————————————————19

B

Baker, H. B., cited —————————————————— 3, 28 

Basommatophora —————————————————— 21, 30 

origin———————————————————————————30 

Bell, W. A., cited——————————————————————6 

Belmont County, Ohio ——————————————————3 

Bensonella —————————————————————— 22

Bernicia praecursor ———————————————————28

Big Creek Shale Member ————————————————— 4

bigsbii. Anthracopupa ——————————————————9

Dendropupa ——————————————————— 9

Pupa ———— 2, 3. 6, 9. 19, 29, 31, 34; pis. 2, 8

bigsbyi, Pupa ———————————————————— 9

Birds, origin ————————————————————————30 

Bostryx————————————————————— 12

Boysidia——————————————————————— 22

Bradley, F. H., cited———————————————————12 

quoted —————————————————————————— 3 

Breezy Hill Limestone Member,

Cabaniss Formation ———————————————4

britannica, Anthracopupa ————————————— 23, 26

Buliminus labrosus labrosus —————————————13

Bulimulidae———————————————————— 12, 13, 21

Austro-Pacific————————————————————— 12

Neotropical ——————————————————————12

Cabaniss Formation, Breezy Hill Limestone Member 

Kansas ——————————————————————— 4 

Carbondale Formation, Herrin Coal Member ——— 4 

Kewanee Group ————————————————————— 4 

St. David Limestone Member———————————28 

Spring Lake Coal Member —————————————4 

Carboniferous, Coal Measures, Little Vermillion

River, Vermillion Co., Indiana - ——————— 4 

Carbonispira——————————————————— 28

scotica ———————————————————— 28

Carpenter, P. P., cited ——————————————————— 7 

Carroll, R. L., cited————————————————————32 

Carychium——————————————————— 21, 30

Casstdula———————————————————— 21

Ceres ————————————————————————— 20

v ^

Cesky Brod, Prague, Czechoslovakia ————————27

Charopa —————————————————— 7

Charopidae ————————————————————————— 8 

Chondrina —————————————————————22

Cionella lubrica ————————————————32

Clams, freshwater —————————————————————30 

Clausiliidae———————————————————————— 12 

Coal beds, fossils —————————————————————3 

Coal-Measures, South Joggins, Nova Scotia ————3 

Coal mine, Little Vermilion River, Ind————————4 

Coal mine, Vermillion Co., Ind————————————25 

Coal No. 4, Summum coal—————————————— 4, 5 

Coal No. 6, Herrin coal——————————————————5 

Illinois section —————————————————————4 

Conemaugh Group ————————————————————— 5 

(Conulus) priscus, Zonites ————————— 6, 7, 9

Cooke, C. M., quoted ———————————————————34 

Cracraft, Joel, quoted ——————————————————30

Page 

cronkhitei, Discus ———————————————32

Cumberland Group————————————————————— 6 

Cyane ————————————————————— 20

Cyclophoridae ————————————————————— 3, 20 

Cyclophoroid taxa ————————————————— 11, 12 

Cyclothem —————————————————————————— 3 

cylindrella, Fernandezia —————————— 21, 22

D

D'Aiguillon, Gaspe Peninsula, Quebec, Canada—28

Dawson, J. W., cited ——————————— 3, 6, 7, 14, 27

quoted ——————————————————————— 6, 10

Dawsonella———————————3, 24, 25, 29, 31, 34

meeki———————————————— 2, 3, 4, 25, ; pi. 10

Dawsonellidae ——————————————————————— 3

Dawson ellinae———————————————————————25

Dendropupa — 2, 3, 6, 11, 12, 13, 26, 27, 28, 29, 31, 32

bigsbii —————————————————————————— 9

Canada ———————————————————————— 33

France—————————————————————————33

geographic range ——————————————— 27, 28

growth pattern————————————————————13

grandaeva ————————————————— 14, 29

grandaevus —————————————————— 14

Poland—————————————————————————33 

primaeva —————————————— 14, 27, 29

ten uistriata ———————————————— Q

thanatocoenose ———————————————————32

uetusta ———— 3, 6, 10, 15, 24, 26, 27, 29; pis. 4, 8

mode of preservation —————————————18

postapical sculpture——————————————10

tenuistriata ————————————————————10

uetusta ———————————————————————10

walchiarum —————————————————14, 26, 27

zarecznyi—————————————— 11, 14, 27

Des Moinesian Series, Illinois —————————————4 

Dendropupinae ——————————————————3, 13, 29 

diminuta, Naticopsis——————————3, 5, 15; pi. 1

Diotocardia ————————————————————————— 29 

Diplommatinoid taxa ——————————————————11 

Discidae——————————————————— 8, 9, 29, 31, 34 

Holarctic ——————————————————————— 34 

Discus ———————————————————— 7, 9

cronkhitei—————————————————— 32

Draparnaudia—————————————————— 13

Dunkard basin——————————————————————— 3

gastropods ——————————————————————— 34

swamp-lake complex —————————————————5

Dunkard beds———————————————————————31

Dunkard Group —————————————————— 5, 24, 36

Greene Formation——————————————————— 5

Uniontown Formation ———————————————35

Washington Formation ———————————————5

Waynesburg Formation ——————————————35

dunkardana, Anthracopupa ————————3, 5, 15; pi. 1

E

Ecole de Mines———————————————————————26

Eggenberg, Austria ———————————————————27

Ellobiidae—————————————————3, 20, 21, 23, 30

freshwater ——————————————————————— 30

Holocene————————————————————————21

Ellobium—————————————-—-—-—— 21

Endodontidae———————————————————8, 20, 34 

Endodontoid, sculpture——————————————————— 8

39
40 INDEX

Page

Enidae———————————————— 12, 13, 21, 29, 31, 34

distribution——————————————————— 13, 34

fossil record, Nova Scotia —————————————34

growth pattern————————————————————13

Brian plant beds —————————————————————— 3

Estrada Nova Formation, Parana basin, Brazil—28

Eurypterids ————————————————————————— 28

F

Fauxulus ————————————————————— 22

Fern Ledges, Lancaster, St. John County,

New Brunswick ————————————————— 3

Little River Group ——————————————————7

Fern trees, Joggins Formation ————————————32

Fernandezia cylindrella——————————— 21, 22

Fire clay, Petty's Ford, Little Vermilion River ———4

Fish remains ———————————————————————28

Flammulina ——————————————————————————9

Floras, Cathaysean ———————————————————33

Euramerican, evolutionary trends ———————33

Gondwana ——————————————————————— 33

Forests, tropical—————————————————————— 32

Fossils, Dunkard basin——————————————————3

Kewanee Group ———————————————————— 3

land snails———————————————————————27

method of analysis——————————————————2

ontogeny ———————————————————————— 2

plants —————————————————————————— 6

spores ——————————————————————————— 6

structure————————————————————————2

Westphalian B, Canada ——————————————— 3

G

Gaspe Peninsula, Canada ——————————————— 28 

Gastrocopta pentodon —————————————— 10

tappamana ———————————————————————10 

Gastropoda —————————————————————————29 

Gastropods, Dunkard basin ———————————————34

freshwater—————————————————— ——— 28 

Paleozoic ————————————————————— 28

terrestrial ——————————————————————— 28

Illinois———————————————————————5

Gibbulina——————————————————————— 22

Gilboy Sandstone Member, Monongahela

Formation — ————————————————————— 

Glaciation, North America ——————————————— 

gondu-anica, Hydrobia —————————————————— 

gossei, Micmceramus —————————————

grandaeva, Dendropupa ——————————————— 14,

Strophella ——————————————— p l

Strophites ———————————————————— 3, 7, 

grandacvus, Dendropupa —————————————————

Strophel/a ———————————————— 14; pi

Strophites ——————————————— ;{, i;{,

Granopupa —————————————————————————— 22 

Greene Formation, Dunkard Group —————————— 5

Nineveh Limestone Member————————— 16, 34 

Gurley Collection —————————————————————25 

Gylnttrachela —————————————————————— 22

H

Hacquebard, P. A., cited——————————————————ti 

Hadrodon——————————————————————;\()

Harry, H. W., cited ————————————————————— 21

Hawaii ————————————————————————————34

Hehcma ————————————————————————————33

Helicinidae ——————————3, 20. 25, 26. 29, 31, 32, 34

distribution, disjunct—————————————————33

tropical —————————————————————————— 33

Helicoid, sculpture —————————————————————iS

Henderson, Junius, cited —————————————————3

Hendcrsoma ——————————————— ;i;i, ;14

Herrin coal, Coal No. 6——————————————————5 

Herrin Coal Member. Carbondale Formation ———— 4 

Hodges Creek, III ——————————————————————5 

Macoupin Co., Ill ——————————————————25 

Holopoda————————————————————————— 1, 31 

Holopodopi's ————————————————————,'i!

Holospira ———————————-——————— 12, 13

Hvdrobia —————————————————————— 2H

Page 

gondivanica—————————————————— 28

Hydrobiidae ———————————————————————— 30

Hypselostoma————————————————22

Illinois basin, limestones, freshwater ————————— 5 

Illinois coal basin ————————————————————— 3 

Introduction ————————————————————————— / 

isoclina, Protornatellina——————————————— 34

Jasper County, 111——————————————————————5 

Joggins area, Nova Scotia ———————————3, 31, 32 

Joggins Formation, tree stumps———————————31 

Journal de Conchyliologie———————————————26 

Juan Fernandez —————————————————————34 

Jurassic-Cretaceous boundary —————————————30

K

Kanawha Formation ———————————————————— 7

Karnovice Limestone, algal bed ———————————27

isotopic composition —————————————————27

lower limestone ———————————————————27

Poland—————————————————————————27

reed bed ————————————————————————27

Karroo Beds, Southern Rhodesia ———————————28

Keele beds, Hagley Park, Birmingham, England—26

Kewanee Group, Carbondale Formation ———————4

Knight, J. B., cited————————————— 6, 10, 11, 25

Kobelt, Wilhelm, cited——————————————————13

Kondo, Yoshio, quoted ——————————————————34

L

labrosus, Buliminus labrosus———

labrosus, Buliminus——————

Lacustrine deposits. Dunkard basin

Lamellidea pusilla

Land snail, England———————————————————26

German—————————————————————————26

Paleozoic, French ———————————————————26

Land snails, apical sculpture ——————————————— 8

biogeographic significance ———————————— 33

carboniferous ————————————————————— 1

deposition ———————————————————— 31, 32

deposits, Mexico ———————————————————32

Texas———————————————————————32

distribution ———————————————————— 33, 34

diversity ———————————————————————— 34

Dunkard basin————————————————————— 6

Dunkard Group ———————————— ——————— 2

ecological significance ——————————————— 31

Fern Ledges, St. John, New Brunswick ————— 6

flood plain, dispersal ————————————————— 32

fossil record——————————————————————— 31

Pennsylvaman—————————————————— 1

fossils, Gondwana areas ——————————————33

geographic distribution ———————————————34

Hudson Bay ——————————————————————32

Illinois——————————————————————————— 3

Illinois basin ——————————————————————6

Indiana—————————————————————————— 3

Joggins. Nova Scotia —————————————— 6, 32

Karniovice. Krakow, Poland————————————27

Monongahela Group —————————————————2

New Brunswick ————————————————————— fi

Nova Scotia ——————————————————————— 6

Ohio ———————————————————————————— 5

Paleozoic ————————————————————— 29, 32

check lists —————————————————————3

ecology —————————————————————————31

local occurrence —————————————————— 3

Indiana——————————————————————— 4

Old World ————————————————————27

stratigraphic setting ——————————————— 3

systematic list ——————————————————29

Pennsylvania —————————————————————— 5

Pennsylvanian—————————————————————29

Permian —————————————————————————29

Petty's Ford, Little Vermilion River ———————4

Polynesia ————————————————————————34

Page 

Land snails—Continued

postapical sculpture————————————————— 8 

prosobranch, freshwater —————————————33 

pupillid, North America——————————————31 

radiation, explosive —————————————————32 

St. John, New Brunswick —————————————6 

Vermillion County, Ind ———————————————4 

Visean—————————————————————————33 

West Virginia ————————————————————— 5 

Westphalian B ——————————————————————— 33 

Launa ——————————————————————— 22

leai, Stenotrema ————————————————— 25

Licharev, Boris, quoted—————————————————27

Limestone, concretionary, Illinois section —————— 4

marine——————————————————————————5

organ pipe ——————————————————————— 27

Vermillion County, Ind ———————————————4

Limestone deposits, Belmont County, Ohio —————5

Limestones, freshwater—————————————————32

Illinois basin —————————————————————5

Jasper County, 111————————————————5

lacustrine ———————————————————————— 6

Limopsis-————————-——-————————— 30

hmpida, Vitrina-———-——————--———32

Lipiarski, Ireneusz, cited ————————————————27 

Lissoacme ———————--—--———————— 12

Little Captina Limestone Member,

limestone, rubbly —————————————————5

Monongahela Formation———5, 23, 24, 31, 34, 35

thin sections —————————————————————16

Little River Group, Fern Ledges ———————————— 7

Fern Ledges. St. John, New Brunswick ————— 6

Little Vermilion River, land snails ——————————4

Locality register, Field Museum

of Natural History————————————————— 35

U.S. Geological Survey——————————————— 34

Monongahela-Dunkard Gastropods———————34

Lower Beaufort beds, Weltevrede, South Africa —28

Lower Washington Limestone ————————————— 5

Lower Washington Limestone Member ————— 6, 18

thin sections —————————————————————16

Washington Formation —————————16, 24, 35

Loxoncma parva————————————— ;i, 5, 24;pl.l

lubrica, Cionella ————————————————— 32

Lucidella --————-———————-———— 33

Lyell, Charles————————————————————————— 6 

Lymnaeidae —————————————————————————30 

Lyrodes---—--—-—--—-——————---——-- 28

M

McFarland, H. B., cited ——————————————— 4, 25 

McLeansboro Group, Illinois ———————————————4

Mattoon Formation —————————————————5 

Macoupin, 111., freshwater beds—————————————25

Jasper Co., Ill—————————————————————25 

Macoupin County, 111 ———————————————————5 

malachoi'i. Omphaluptychia —————————— 27

Mangrove swamp ————————————————————— 30 

Mansfelder Schichten, Halle, West Germany———26 

Matthew, G. F., cited ——————————————————— 6

quoted———————————————————————— 7, 14 

Mattoon Formation, limestone, marine ————————5

McLeansboro Group —————————————————5

Reisner Limestone Member—————————————5 

Matunpupa————————————————:}, 20, 26

I'crrnilwni'nsis ——————————————24

mceki. Anomphalus———————————————25

Dau'sonclla —————————— 2, 3, 4, 25: pi. 10

Mesauricu/stra --———————--———-———— 30

Meswhilina——————————————————30

Mesogastropods———————————————————————12 

Mcsoncritina————————————————— 29, 30

Mesurethra ——————————————————————— 1, 31 

Micrwcramus———————————————— 10, 12

gossri—————————————————————— 12

Micronesia——————————————————————————34 

minutistiioniim. Punctum————————————— 32

Modiomorpha————————————————— 28

Mollusks, freshwater————————————————— 30, 31 

Monongahela-Dunkard Gastropods, locality

registers———————————————————————34
INDEX 41

Page 

Monongahela Formation,

Gilboy Sandstone Member —————————35 

Little Captina Limestone Member——— 5, 23, 24,

31, 34, 35

Monongahela Group ———————————— 5, 24, 35, 36 

Uniontown Formation ———————————————35 

Monsoon areas———————————————————————32 

Morrison Formation, North America ————— 29, 30 

morsel. Vertigo ————————————————— 19

Museum National d'Histoire Naturel, Paris ———— 26 

Myalinidae ———————————————————————— 30

N

Naticopsis dimmuta —————————3, 5, 15; pi. 1

Neogastropods ———————————————————————12

Neritacea ——————————————————————————29

freshwater———————————————————————29

Great Britain —————————————————————29

marine deposits ———————————————————29

terrestrial———————————————————————29

Newell, N.D., cited ————————————————————28

Ninevah limestone ————————————————————35

Nineveh Limestone Member, Greene Formation 16, 34

Nipa palm swamp ————————————————————30

notata, Triodopsis (Xolotrema) ————————— 25

o

oblongus, Tornatellides ———————————— 23

Odontocyclas —————————————————— 22

ohioensis, Anthracoupa ———3, 4, 5, ;5, 20, 23, 24,

25, 26, 27, 29; pis. 1, 5, 6, 7, 9

Oligyra————————————————— 26, 32, 33

orbiculata tropica ————————————— 26

Omphaloptychia malachovi ———————————27

oncospira, Tornatellides—————————————— 21

orbiculata tropica, Oligyra ———————————————26 

Orobophana —————————————————— 26

Orthuretha —————————————————————— 29, 31 

oswaldi, Tornatellides —————————————————— 21 

Otinidae———————————————————————————30

Parana basin, Brazil———————————————————28 

Partulacea ————————————————————————— 29 

Partulidae —————————————————————————34 

parva, Anthracopupa ——————— 23, 24, 29; pi. 1

Loxonema —————————————;}, 5, 24; pi. 1

Peck, J. H., cited——————————————————— 4, 25 

pentodon, Gastrocopta —————————————————10 

Petersburg Formation, Indiana—————————————4 

Petty's Ford, Little Vermilion River, 111———————25 

Phyletic significance ———————————————————29 

Physidae ——————————————————————————30 

picta, Anguispira ———————————— 8, 9; pi. 3

Pilshury, H. A., cited——————————— 3, 7, 8, 21, 28

quoted—————————————————————— 12, 20 

Pineria —————————————————————— \2

Planogyra —————————————————————————— 8 

Planorbidae————————————————————————30 

Plant bed No. 2, Fern ledges, St. John,

New Brunswick————————————————— 6, 7

Plant beds, below coal No. 6, Petty's Ford, Vermilion

River, 111. ——————————————————————3

Brian —————————————————————————— 3

St. John, New Brunswick —————————————3 

Plants, arborescent ———————————————————33

continuous leaf-shedding ————————————— 33

laminate foliage ———————————————————33 

Pleasant Grove, Belmont County, Ohio ——————— 5

Belmont County, Ohio, Permian rocks ————— 3 

Pleuropoma ———————————————————————— 26 

Polynesia——————————————————————————34 

Pomatiasidae, West Indian ———————————————11 

Poole, E. G., cited —————————————————————26 

Potamopyrgus ——————————————————— 28

praecursor, Bernicia———————————————————28 

primaeva, Dendropupa——————————— 14, 27, 29

Pupa ————————————————— 2, 3, 7, 14

priscus, Archaeozonites —————————————— 7

Protodiscus —————————————— 2, 29 pis. 2, 8

priscus—Continued Page 

Pyramidula —————————————————— 7

Zonites ——————,——————————— 3, 7, 8, 9

(Conulus)————————————— 6, 7, 9

Proserpina ———————————————————— 20

Prosobranchia ———————————————————————29 

Prosperinella ——————————————————— 20

Proserpininae ——————————————————————— 20

Prosobranch, thiarid ———————————————————28

Protodiscus ———————————————— 2, 10, 28, 31, 34

priscus————————————————2, 29; pis. 2, 8

Protornatellina isoclina ————————————— 34

Pruvost, Pierre, cited———————————————————26 

Pseudhelicinia——————————————— 26, 33

Psuedosigmurethrous features ————————————— 31 

Pterodactyl ———————————————————————— 33 

Pulmonata—————————————————————————29 

Pulmonate——————————————————————— 7, 21 

Punctidae——————————————————————————8 

Punctum ————————————————————— 8

minutissionum———————————————32

Pupa bigsbii———————————————2, 3, 6, 9, 19,

29, 31, 34; pis. 2, 8 

bigsbyi ——————————————————— 9

primaeva —————————————2, 3, 7, 14

vermilionensis ———————— 3, 4, 5, 20, 24; pi. 1

vetusta ———————————————————3, 6, 10, 13

tenuistriata ———————————————————10

Pupilla —————————————————————————-— 10

Pupillacea ———————————————— 10, 12, 29, 31, 34

Pupillidae———————————————————————— 3, 20

distribution —————————————————————— 34

recent——————————————————————————19

Pupilloid, growth pattern————————————————21

Pupilloid, sculpture ————————————————————8

pusilla, Lamellidea ——————————————— 23

Pyramidula ————————————————————————— 7 

priscus——————————————————— 7

Pythia———————————————————————— 21

R

Radiodiscus———————————————————— 8

Rain forest areas—————————————————————32 

Redpath Museum, collection———————————————6 

References cited—————————————————————— 36 

Reisner Limestone Member, Mattoon Formation — 5 

rothenburgensis. Anthracopupa————— 23, 26, 27

Runnegar, Bruce, cited—————————————————28 

Russell, L. S., quoted———————————————————28

St. David Formation, 111 ————————————— 25

St. David Limestone Member, Carbondale

Formation—————————————————————28

Saone-et-Loire———————————————————————26

Schasicheila ———————————————————————— 26

Schroyer, C. R., quoted —————————————————— 5

scotica, Carbonispira———————————————————28

Scott's Cut, Belmont County. Ohio —————————35

Scremerston Coal Group —————————————————28

Scremerston, Northumberland, England —————— 28

Sigillaria —————————————————————————— 6

Sigmurethra————————————————————— 29, 31

Slugs ————————————————————————————— 1

Snail, planorbiform, Madagascar———————————28

Snails, freshwater—————————————————— 30, 31

freshwater. Paleozoic————————————————— 28

hydrobiid, freshwater ————————————————28

Paleozoic, nonmarine ———————————————26

Sukhona-Mezen Rivers area, Vologda region,

Moscow———————————————————————27 

terrestrial ——————————————————————— 31 

Spirorbis———————————————————— 5, 24, 25, 27 

Spring Lake Coal Member, Carbondale Formation-4 

Staesche, Karl, cited ———————————————————27 

Stauffer, C. R., quoted ——————————————————— 5 

Stenotrema lea/ ——————————————————————25 

Stratigraphic correlations, North America

and Europe —————————————————————3 

Striatura————————————————————— 8

Strophella ——————————————— :i, 13, 14, 15

Strophelta—Continued Page 

grandaeva ——————————————————pi. 1

grandaevus ————————————— 14; pi. 1

Strophites ——————————————————— 3, 13

grandaeva ——————————————— 3, 7, 15

grandaevus———————————————3, 13, 14

sturgeoni, Anthracopupa —————— 2, 23, 26, 29; pi. 9 

Stylommatophora ———————————————— 29, 30, 31 

origin——————————————————————————30 

Subulinidae—————————————————————— 12, 21 

Sukhona-Mezen Rivers area, Vologda region,

Moscow———————————————————————27 

Summum coal, Coal No. 4—————————————— 4, 5 

Systellommatophora ——————————————————— 1 

Systematic summary———————————————————28 

Systematic work, earlier—————————————————3 

Systenostoma———————————————————22

tappaniana, Gastrocopta - 

Taxa, analysis ———————

classification ————— 

tenuistriata, DendropupaDendropupa vetusta—

Pupa vetusta-

—— 10

——— 7

——— 7

——— 6

——10

——10

Terrestrial strata, New Brunswick———————————3 

Top Hosie Limestone———————————————————28 

Tornatellides oblongus ————————————— 23

oncospira————————————————— 21

oswaldi —————————————————— 21

Tornatellinidae ———-———3, 10, 12, 20, 23, 29, 31, 34

distribution ——————————————————————34

speciation centers ——————————————————34

Tornatellinoid, growth pattern ————————————21

Tortacella———————————————————— 30

Travertine ————————————————————————— 27 

Tree stumps —————————————————————— 6, 31 

Triodopsis (Xolotrema} notata —————————————25 

tropica, Oligyra orbiculata ——————————26

u

Ulpia —————————————————— 22

Unio—————————————————————————————30

Unionid precursors———————————————————— 30

Unionidae ————————————————————————— 30

Uniontown Formation —————————————————35

Dunkard Group ———————————————————35

Monongahela Group ————————————————35

Uniontown Limestone Member——————— 23, 24

Uniontown Limestone Member,

Uniontown Formation ——————————— 23, 24 

Upper Washington Limestone Member,

Washington Formation —————————————35

Uptown borehole, Oxfordshire, England —————— 26

Urocoptid stock, Antillean ———————————— 12, 13

Urocoptidae——————————————————— 3, 12, 21,29

convergences —————————————————————12

distribution ——————————————————————13

growth pattern————————————————————12

origin———————————————————————————13

V

Valvatidae—————————————————————————30 

ventricosa. Vertigo———————————————32

Vermilion County, 111 ———————————————————3

vermilionensis, Anthracopupa- 23, 24, 25, 27, 29; pl.l

Ma turipupa ————————————————— 24

Pupa —————————————— 3, 4, 5, 20, 24; pi. 1

Vermilionville Sandstone Member———————————4

Vertigo ————————————————————————————10

morsei —————————————————————————19

ventricosa ———————————————————————32

Vetulonaia—————————————————————————30

vetusta, Dendropupa ———————————3, fi, 10, 15, 24,

2fi, 27, 29; pis. 4, 8

Dendropupa vetusta——————————— 10

Pupa —————————————————————3, 6, 10, 13

tenuistrata, Dendropupa —————————————10

Pupa —————————————————— 10

vetusta. Dendropupa —————————————————10
42 INDEX

Page

Vitrina limpida ——————————————————————32 

Viviparidae—————————————————————————30 

Vologda region, Russia—————————————————27

w

walchiarum, Dendropupa ————————— 14, 26, 27

Waldemaria—————————————————— 26, 34

Walden, Barbara, cited ————————————————— 21

Washington Formation, Dunkard Group———————5

lower limestone member———————————————5

Lower Washington Limestone Member-16, 24, 35

Page

Washington Formation—Continued

Upper Washington Limestone Member ————35

Waterfowl, Australian——————————————————30

European —————————————————————30

Waynesburg Formation, Dunkard Group —————35

Weber, Jon, cited—————————————————————27 

Weller, J. M., cited ———————————————————4 

Weltevrede, South Africa ———————————————— 28 

Wenz, Wilhelm, cited ——————————————————— 21 

White, C. A., cited—————————————————————3 

Whitfield, R. P., cited————————————————— 4, 21 

quoted —————————————————————————— 4 

Worm tubes ————————————————————————— 28

Page

X,Y,Z

(Xolotrema) notata, Triodopsis 

Yen, Teng-Chien, cited —————

Zaptychius————————————————-————— 30

zareczyni, Dendropupa —————————————11, 14, 27 

Zilch, Adolph, cited ————————————————— 3, 21 

Zonites —————————————————————— 6

(Conulus) priscus ———————————— 6, 7, 9

priscus—————————————————— 3, 7, 8, 9

Zonitidae ——————————————————————————— 3 

Zonitoid, sculpture —————————————————————8
PLATES 1-10

Contact photographs of the plates in this report are available at cost, from U.S. Geological 

Survey Library, Federal Center, Denver, Colo. 80225
PLATE 1

[Figs. 1-3, 5-22 are X 15; fig. 4 is X 5]

Figures 1-3, 8-10, 13, 14, 18, 19. Anthracopupa ohioensis Whitfield, 1881, (p. 15).

1, 2. Side view of lectotype of Naticopsis (?) diminuta, OSU (Ohio State University) 15170, probably the original of pi. 11, fig. 23 of Stauffer and Schroyer 

(1920). 3. Oblique side view of lectoparatype of AT.(?) diminuta, OSU 15169, 

probably the original of pi. 11, fig. 22 of Stauffer and Schroyer (1920). Both 

specimens from "the shales at the base of the Lower Washington Limestone, 

one-half mile south of Pleasant Grove, Belmont County, Ohio" (Stauffer and 

Schroyer, 1920, p. 145). 8, 9. Oblique apical and side view of lectoparatype of 

A(?) dunkardana Stauffer and Schroyer, 1920 (p. 15), OSU 15163, possibly 

the original of pi. 11, fig. 14 of Stauffer and Schroyer (1920). 10. Basal view of 

lectoparatype steinkern of A. (?) dunkardana, OSU 15162. 13, 14, 18, 19. 

Holotype, AMNH (American Museum of Natural History) 8487/1. From 

"Marietta, Ohio" (Whitfield, 1881, p. 126). 13. Side view. 14. Oblique apical 

view. 18. Oblique basal view, showing the thickened apertural lip. 19. Apertural view. 

5-7. Anthracopupa vermilionensis (Bradley) 1872, p. 24.

5. Apertural view of Pupa uermilionensis Bradley, 1872, USNM (United 

States National Museum) 242,698. 6. Side view, USNM 242,699. 7. Side view 

with aperture broken back, USNM 242,700. All specimens out of USNM lot 

27,600 labeled "Upper Carboniferous (Coal No. 6), four miles below Georgetown, Illinois."

4. Strophella grandaeua (Dawson, 1880) (p. 14). Side view of holotype Strophella grandaevus Dawson (1880) with light at an extremely low angle to 

emphasize the ribs, Redpath Museum 2384, from "plant-beds of St. John, 

New Brunswick" (Dawson, 1880, p. 413).

11-12. Anthracopupa^) dunkardana Stauffer and Schroyer, 1920 (p. 15). Side and 

apertural views of lectotype of A(?) dunkardana, OSU 15161, probably the 

original of pi. 11, figs. 12 and 13 of Stauffer and Schroyer (1920). All specimens from the same locality as specimens shown in fig. 1, this plate.

15-17, 20-22. ?Anthracopupa parua (Stauffer and Schroyer) 1920 (p. 24). 15, 20. Apical 

and apertural views of lectotype LoxonemaC!) parua Stauffer and Schroyer, 

1920, OSU 15165. This is the original of pi. 11, fig. 19 of Stauffer and 

Schroyer (1920). 16,17. Slightly oblique side view and oblique apical view of 

lectoparatype LoxonemaC?) parua, OSU 15166. This is the original of pi. 11, 

fig. 21 of Stauffer and Schroyer (1920). 21. Side view of broken lectoparatype 

LoxonemaC?) parva Stauffer and Schroyer, 1920, OSU 15167. This is the 

original of pi. 11, fig. 20 of Stauffer and Schroyer (1920). 22. Side view of 

largest specimen, Loxonema(fl) parua, lectoparatype OSU 15168; the original of pi. 11, figs. 17 and 18, Stauffer and Schroyer (1920); in those figures, 

the aperture was restored. All specimens from the same locality as specimens 

shown in figure 1, this plate.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1072 PLATE 1

STROPHELLA, ANTHRACOPUPA (?), ANTHRACOPUPA, AND ? ANTHRACOPUPA
PLATE 2

FIGURES 1-5. Protodiscus prisons (Carpenter, 1867) (p. 7).

1. X31.6. Holotype. Coal Measures, Nova Scotia. Redpath Museum 3116. Original of Dawson's 

1867 fig. b and 1880 fig. 10. 2-5. Paratype. Coal Measures, Nova Scotia. Redpath Museum 

3116b. 1. X31.6. Oblique apical view of entire shell, partly crushed. 2. X98. Apical and early 

postapical sculpture. 3. X92. Low-angle view of sculpture, apical left, postapical right. 4. X184. 

Detail of apical sculpture. 5. X475. Detail of spiral postapical sculpture near the suture. 

6. Pupa bigsbii Dawson, 1880 (p. 9). Lectotype. Erect trees of Group XV, Coal Measures, South 

Joggins, Nova Scotia. Redpath Museum 3122. Apertural view of entire shell. X42.6.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1072 PLATE 2

PROTODISCUS AND PUPA
PLATE 3

FIGURES 1-6. Anguispira picta (Clapp, 1920) (p. 8). Buck Creek Cove, south of Sherwood, Franklin County,

Tennessee. Glenn Goodfriend. Sept. 6, 1974. FMNH 171138.

1. X72. Apical boundary area. 2. X229. Detail of oblique microsculpture at end of apex. 3. 

X22.6. Early whorls of juvenile shell. 4. X112. Midapical area having radial elements well 

developed, but oblique microelements absent to weakly developed. 5. XI12. Apical transition between microsculpture only, and radial elements present. 6. X228. Detail of oblique 

microsculpture.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1072 PLATE 3

ANGUISPIRA
PLATE 4

FIGURES 1-6. Dendropupa vetusta (Dawson, 1855) (p. 10).

1. X21.8. Coal Measures, Joggins, Nova Scotia. Redpath Museum 15.011. Partly embedded shell 

having 5'/4 whorls preserved. 2. X116. Redpath Museum 15.011. Details of apical (right) and 

first postapical whorl sculpture. 3. X174. Redpath Museum 15.007. Suture on spire of a shell 

fragment. 4. X571. Redpath Museum 15.011. Fracture through fossil showing recrystallization. 5. X2,200. Redpath Museum 15.011. Detail of recrystallization. 6. X79. Redpath Museum 

15.012. Detail of body whorl behind expanded lip.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1072 PLATE 4
PLATE 5

FIGURES 1-6. Anthracopupa ohioensis Whitfield, 1881 (p. 15). "Lower Washington Limestone Member," 

Washington Formation, Ayers limestone quarry in SW J/4 sec. 19, Pease Twp., Belmont 

County, Ohio. USGS 24855-PC (USNM 242, 701).

1. X18.9. Apertural view. Entire shell, surface sculpture eroded. 2. X49. Gerontic aperture. 3. 

X89. Detail of right part of aperture. 4. X178. Parietal barrier, showing crevice left by dissolved periostracum. 5. X872. Break on outer surface of parietal barrier. 6. X4,300. Microcrystals on surface of parietal wall and barrier.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1072 PLATE 5

ANTHRACOPUPA
PLATE 6

FIGURES 1-6. Anthracopupa ohioensis Whitfield, 1881 (p. 15). "Lower Washington Limestone Member," 

Washington Formation, Ayers limestone quarry in SW J/4 sec. 19, Pease Twp., Belmont 

County, Ohio. Juvenile specimens. USGS 24855-PC.

1. X51. Sculpture on shell surface. USNM 242,702. 2. X205. Apical (upper left) and postapical 

sculpture. 3. X64. Juvenile aperture. 4. X89. Detail of aperture. 5. X33. A narrower juvenile 

shell. USNM 242,703. 6. X77. Detail of aperture showing bilayered nature of lip at bottom.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1072 PLATE 6

.5

ANTHRACOPUPA
PLATE 7

FIGURES 1-6. Anthracopupa ohioensis Whitfield, 1881 (p. 15).

1-4. "Lower Washington Limestone Member," Washington Formation, Ayers limestone 

quarry in SWVi sec. 19, Pease Twp., Belmont County, Ohio. USGS 24855-PC. (Barrier lost; 

remainder of specimen, USNM 242,704.) 1. X138. Excised parietal barrier, anterior margin 

at left. 2. XI,326. Side of barrier near posterior end showing change in size of microdenticles 

from side to top of barrier. 3. X 1,695. Microprojections on top of parietal barrier. 4. X9,764. 

Details of individual microprojections on barrier top.

5-6. Basal part of Little Captina Limestone Member, Monongahela Formation, ravine on 

north side of Deep Run Road, extreme west sec. 26, Pease Twp., Belmont County, Ohio. 

USGS 24836-PC. (Specimen lost.) 5. X171. Anterior end and upper edge of columellar barrier. 6. X888. Microprojections on barrier (right) and matrix crystals.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1072 PLATE 7

ANTHRACOPUPA
PLATE 8

FIGURES 1, 2. Protodiscus priscus (Carpenter, 1867) (p. 7). Coal Measures, South Joggins, Nova Scotia. Paratype. Redpath Museum 3116a. Basal view of shell. 

1. X21.3. View from basal angle. 2. X22.2. View from apical angle.

3, 4. Anthracopupa sp. (p. 10). Paratype of Pupa bigsbii Dawson, 1880. Redpath Museum 3122a. 

Coal Measures, South Joggins, Nova Scotia. 3. X42.5. Side view. Note remnant radial ribbing 

at upper left. 4. X58. View of worn side of apex.

5. Dendropupa vetusta (Dawson, 1885) (p. 10). Coal Measures, South Joggins, Nova Scotia. Redpath Museum 15.010. X50. Isolated columellar fragment showing the two raised ridgelike barriers on surface.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1072 PLATE 8

PROTODISCUS, ANTHRACOPUPA, AND DENDROPUPA
PLATE 9

FIGURES 1-5. Anthracopupa sturgeoni Solem and Yochelson, n. sp. (p. 23). Little Captina Limestone Member, Monongahela Formation, in ravine on north side of Deep Run Road, extreme west sec. 26, 

Pease Twp., Belmont County, Ohio. 

1-3, 5-Holotype. USGS 24836-PC. (USNM 242,705). 

1. X31.5. Apertural view of entire shell. 2. X59. Detail view of aperture. 3. X56. Lateral view of

aperture. 5. XI14. Detail of shell sculpture on spire. 

4. Paratype. USGS 24836-PC. (USNM 242,706). X34.5. Apical fragment. 

6. Anthracopupa ohioensis Whitfield, 1881. (p. 15). Nineveh Limestone Member, Greene Formation, center SWV4 sec. 16, Salem Twp., Monroe County, Ohio. USGS 24827-PC. (USNM 

242,715). X112. Shell sculpture on spire for comparison with sculpture of A. sturgeoni.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1072 PLATE 9

ANTHRACOPUPA
PLATE 10

FIGURES 1-6. Dawsonella meeki (Bradley, 1872) (p. 25). Petty's Ford, Little Vermilion River, below Georgetown, Vermilion County, Illinois FMNH 11673 from the Gurley Collection. 

1. X38.9. Aperture. 2. X42.3. Lateral view of aperture to show basal-columellar projection. 

3. X107. Sculpture on base of shell. 4. X100. Upper surface of shell showing broken layers. 

5. X997. Detail of broken shell layer. 6. X4,985. Detail of crystal pattern in upper broken layer.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1072 PLATE 10

DA WSONELLA