J. Paleont., 83(6), 2009, pp.

938–945
Copyright ’ 2009, The Paleontological Society
0022-3360/09/0083-0938$03.00

LATE PALEOZOIC CONTINENTAL GASTROPODS FROM POLAND:

SYSTEMATIC, EVOLUTIONARY AND PALEOECOLOGICAL APPROACH
EWA STWORZEWICZ,1 JOACHIM SZULC,2 AND BEATA M. POKRYSZKO3
1
Polish Academy of Sciences, Institute of Systematics and Evolution of Animals, Sławkowska 17, 31-016 Cracow, ,stworzewicz@isez.pan.krakow.pl.;
2
Jagiellonian University, Institute of Geological Sciences, Oleandry 2a, 30-063 Cracow, ,joachim.szulc@uj.edu.pl.; and 3Wroclaw University,
Museum of Natural History, Sienkiewicza 21, 50-335 Wrocław, Poland, ,bepok@biol.uni.wroc.pl.

ABSTRACT—Two taxa of the Late Carboniferous and four species of the Early Permian terrestrial snails have been
found in the Late Paleozoic continental molasse sediments of the Upper Silesian-Cracow Upland (Southern Poland).
Discovery of Anthracopupa ohioensis and Protodiscus priscus indicates that, besides in North America, they occurred
also in the European part of the Pangea supercontinent. According to the general sedimentary facies context and the
accompanying floral and faunal assemblages, the gastropods lived in swamp environments, including a topogenous
fen.

INTRODUCTION It is noteworthy that organic remains are rarely found in

of the first terrestrial snails and their differen-
T HE ADVENT
tiation in the earliest stages of their evolution have long
been of great interest to malacologists, this being among the
crucial problems of gastropod evolutionary relationships. The
extreme paucity of the fossil record does not enable a reliable
reconstruction of the early history of the terrestrial malaco-
fauna. The Upper Silesian–Cracow area is among the few
regions with a fossil record of land snails from the Late
Palezoic. The most abundant data come from the Upper
Palezoic rocks of the United States and Canada. The existing
fossil record indicates that at the end of the Paleozoic,
terrestrial snails were already quite diverse. Solem and
Yochelson (1979) assigned them to three out of the six orders
known today and recognised 10 pulmonate and one terrestrial
prosobranch species, whereas Bandel (2002) included the Late
Paleozoic gastropods in a new order, the Procyclophorida.
Based on the previous studies, the total number of terrestrial
Paleozoic snails worldwide does not exceed 25.
To date, the only Polish contribution to these studies was a
description of Dendropupa zarecznyi by Panow (1936) from
the Lower Permian freshwater limestones of Karniowice, near
Cracow. The present study yields many new specimens,
representing some additional species and allowing a re-
evaluation of the previous results of study of the Late
Paleozoic continental gastropods.
Beside the previously known Early Permian malacofauna,
there was no published information on gastropod remains from
other sites from southern Poland. The recent findings of the
gastropods from the Upper Carboniferous terrestrial sediments
(Filipiak and Krawczyński, 1996; Krawczyński et al., 1997)
provided specimens representing probably two snail species.
LOCATION AND MATERIAL
Carboniferous siderite nodules from Sosnowiec (Fig. 1,
locality 1).—The snail-bearing spherosiderite concretions were
found on a spoil-heap of the Pora˛bka–Klimontów coal mine
in Sosnowiec, in the NE. part of the Upper Silesian Mining
Region (50u179N, 19u079E). Their age was estimated as
Westfalian A by means of palynological and macrofloral
analysis (Krawczyński et al., 1997; Pacyna, 2003). The
concretion-bearing rocks also comprise other freshwater and
terrestrial fauna, such as crustaceans, bivalves, insects, and
remains of terrestrial plants of the genera Calamites, Sigillaria,
and Lepidostrobus.
938
spherosiderites. Out of several thousand siderite nodules
collected from spoil-heaps in Sosnowiec, only a few dozen
contained well-preserved fragments of plants and animals
(Krawczyński et al., 1997). Molluscs were found in only four
concretions, one of which contained bivalve remains, while the
remaining ones held three snail imprints (ISEZ-K/93–ISEZ-K/
95) described here for the first time.
Because of the great diversity of spherosiderite fossils, the
site in Sosnowiec is very valuable, especially from a faunistic
viewpoint, ranking beside the two most famous sites with
remains of Carboniferous plants and animals in Mazon Creek
(Illinois, USA) and at Joggins (Nova Scotia, Canada) (Pacyna
and Zdebska, 2002; Hebert and Calder, 2004; Falcon-Land et
al., 2006). Faunal assemblages known from other European
coal deposits are distinctly poorer.
The snails found in three spherosiderite concretions are shell
imprints, and the shell form was reconstructed as moulds
using plastic mass. In only one case was the condition of the
imprint sufficiently good to permit a reconstruction of the
exact shape of the shell and its aperture.
Lower Permian freshwater carbonates – ‘‘Karniowice Trav-
ertine’’ (Fig. 1, locality 2).—In the region of Cracow, Zare˛czny
(1894) found Paleozoic deposits containing gastropod shells as
well as numerous plant remains. These materials originating
from the so-called ‘‘Karniowice travertine’’ served as the basis
for a paper by Panow (1936), containing a description of a new
species, while mention of some other snails from the site
appeared in Solem and Yochelson (1979).
Carbonate deposits in the region of Krzeszowice near
Cracow (50u099N, 19u339E), often collectively referred to as
‘‘Karniowice travertine,’’ are especially interesting among
other Lower Permian sedimentary rocks of the Cracow region.
Recent studies (Szulc and Ćwiz_ewicz, 1989; Ćwiz_ewicz and
Szulc, 1989) showed that the spectrum of carbonate rocks is
much broader than travertine formations sensu stricto; other
rocks include calcareous deposits formed in small stagnant
water bodies (semi-limnic), in streams feeding these ponds,
and calcareous deposits of swamp facies with numerous
calcified remains of vascular plants (Fig. 2). Based on a very
rich assemblage of such plants, Lipiarski (1971) described their
habitat as a topogenous fen. The vascular plant assemblage
includes both Carboniferous and Permian components, which
makes it difficult to unambiguosuly determine the age of the
Karniowice travertine (Raciborski, 1891). The ultimate
 STWORZEWICZ ET AL.—PALEOZOIC GASTROPODS FROM POLAND
FIGURE 1—Location map of the gastropod occurrences. 1, Upper
Carboniferous gastropods; 2, Lower Permian gastropods.
criterion for classifying the travertine as Lower Permian was
the high proportion of new components that were absent from
Carboniferous assemblages (Lipiarski, 1971). Survival of
Carboniferous tropical flora in the generally semi-arid climate
of the early Permian was possible only due to the presence of a
vast system of karstic springs that fed an extensive oasis.
As mentioned above, the first Paleozoic land snails were
found over a hundred years ago by Zare˛czny (1894). Based on
his materials, Panow (1936) described Dendropupa zarecznyi.
He mentioned also several other specimens of other species
collected by him, but these have never been examined or
described. General information on Dendropupa snails from
Karniowice was provided by Solem and Yochelson (1979).
Yochelson visited the site in 1968 and collected a few more
specimens, stored in the National Museum of Natural History
of Smithsonian Institution in Washington.
The material for this study was collected by the authors
from 2004 to 2007. It includes 92 limestone samples with snail
fossils (ISEZ-K/1–ISEZ-K/92), from one to seven per frag-
ment (a total of 170 specimens). Unfortunately, poor
preservation of the specimens often precludes precise species
identification. Only a few specimens show traces of shell
surface sculpture. The collection is kept in the Institute of
Systematics and Evolution of Animals, Polish Academy of
Sciences, Cracow. Additional material comes from Panow’s
collection stored in the Geological Museum PAS, Cracow
(A-I-56/1–A-I-56/27).
All measurements were taken with Nikon measurescope
MM-11, accuracy 0.01 mm.
PALEOGEOGRAPHICAL AND PALEOENVIRONMENTAL CONTEXT
At the end of the Carboniferous, a collision of continental
plates led to the formation of the large supercontinent Pangea
and to origin of the Variscian orogenic belt, with a vast foreland
basin system filled by coal-bearing molasse deposits (Fig. 3).
The continental plate repositioning resulted in changes to
atmospheric circulation and moisture distribution and brought
about floristic changes (Opluštil, 2004). The paleogeographical
position of the Variscan belt imposed tropical climatic
conditions in the late Carboniferous, generally hot and humid.
Wetland terrestrial ecosystems, reconstructed on the basis of
939
FIGURE 2—Scheme of the Lower Permian carbonate facies distribution
of the Karniowice Travertine. 1, Lower Carboniferous basement rocks; 2,
fanglomerates with pyroclastics; 3, spring deposits –travertine sensu
stricto; 4, paleosoils with caliche crusts.
the Joggins Formation deposits, were characterized by a diverse
vegetation comprising pteridosperms, ferns, lycopsids, cordai-
taleans and calamiteans (Falcon-Land et al., 2006). One facies
of these deposits comprises siderite nodules with tetrapod
skeletal material and some other animals. Siderite nodules from
Sosnowiec also contain a rich faunal assemblage, including
crustaceans and molluscs. The plant macrofossils represent
typical Carboniferous plant taxa, such as lycopods, sphenop-
sids, pteridophylls, cordaites and conifers (Pacyna and
Zdebska, 2001, 2002; Pacyna, 2003). The animal and plant
fossils represent two environments—terrestrial and aquatic
(Filipiak and Krawczyński, 1996; Krawczyński et al., 1997).
The plant remains, insects and snails indicate terrestrial
habitats; the crustaceans are freshwater forms, while bivalves
inhabit both fresh and brackish waters.
The calcified floral remnants in the freshwater limestones
from Karniowice include many taxa also recorded from
deposits of the Joggins Formation. These plant assemblages
represent mainly a hygrophilous or nearly-hygrophilous
vegetation type characterictic of Carboniferous and early
Permian peat associations (Lipiarski, 1971). Based on their
stable isotope study, Szulc and Ćwiz_ewicz (1989) pointed out
that the Lower Permian Karniowice freshwater carbonates
were deposited in subtropical conditions comparable to those
of the present-day subtropics. These carbonates are composed
of travertines, lacustrine, and palustrine limestones deposited
in a spring-fed oasis basin. Such conditions were favorable for
a range of animals, including the earliest terrestrial gastro-
pods, which required a wetland environment at the early
stages of their terrestrial life. It is also noteworthy that the
snails discussed occur only in the palustrine facies of the
Karniowice Travertine limetsones.
The high proportion of small snails is a characteristic
feature of the Karniowice assemblage, while the considerably
larger D. zarecznyi is represented by only few specimens. It
is interesting that D. zarecznyi was always found together
with large plant fragments. In contrast, small shells were
accompanied by fine plant fragments. These facts may indicate
either different microhabitat requirements or slightly different
taphonomic modes.
940 JOURNAL OF PALEONTOLOGY, V. 83, NO. 6, 2009
FIGURE 3—Paleogeographic position of the discussed areas. 1, Cracow-
Upper Silesian area; 2, Joggins Formation of Nova Scotia.
SYSTEMATIC PALEONTOLOGY
Phylum MOLLUSCA Linnaeus, 1758
Class GASTROPODA Cuvier, 1797
Order EUPULMONATA Haszprunar and Huber, 1990
Superfamily ELLOBIOIDEA L. Pfeiffer, 1854
Family ANTHRACOPUPIDAE Wenz, 1938
Genus ANTHRACOPUPA Whitfield, 1881
ANTHRACOPUPA OHIOENSIS Whitfield, 1881
Figure 4.1–4.4
Anthracopupa ohioensis Whitfield, 1881, p. 126, figs. 1–4.
Material examined.—Three specimens ISEZ-K/11, ISEZ-K/
54, ISEZ-K/30 (only the latter in relatively good condition).
Dimensions (in mm).—Height: 4.40, height of last whorl:
3.15; Width: 3.03; Number of whorls ca. 4.5.
Description.—Best-preserved shell of ovoid shape, consist-
ing of 4.5 moderately convex whorls separated by rather
shallow suture. Rounded aperture filled with calcite sediment,
difficult to remove, hence no apertural teeth visible. In another
specimen, due to the body whorl being partly crushed,
something like a columellar tooth can be seen inside
(Fig. 4.2). Similar structure also visible in the polished section
of rock piece with Anthracopupa shell (Fig. 4.4). Aperture
margin broken so that the outer lip thickening is difficult to
define. The surface sculpture also invisible.
Discussion.—The specimens from Karniowice are strikingly
similar to the shorter and more tumid shells of A. ohioensis,
presented by Solem and Yochelson (1979, pl. 1, fig. 19), in
having the same shell shape and form of the aperture
(Fig. 4.3); however the latter are generally somewhat smaller.
ANTHRACOPUPA BRITANNICA Cox, 1926
Figure 4.5–4.7
Anthracopupa britannica Cox, 1926, p. 407, figs. 1a, 1b, 2a, 2b.
Material examined.—five specimens in relatively good
condition (ISEZ-K/71, ISEZ-K/55, ISEZ-K/1a–c) and 115
fragments of shells.
Dimensions (in mm).—Height: 3.36, 3.57, 3.62, 3.90, 4.15;
Width: 1.30, 1.45, 1.55, 1.75, 1.75; Number of whorls 5–6.
Description.—Shell rather slender with moderately con-
vex whorls. Apertures of all specimens damaged, hence the
proper aperture shape and character of outer lip unknown.
In several shells a small parietal tooth is more or less
visible (Fig. 4.6), but only one shell has a structure
resembling a columellar tooth (Fig. 4.5). It can be
assumed that the surface sculpture of the teleoconch
whorls was lost during fossilization, because a very fine
radial striation is still visible in places. Protoconch surface
unknown.
Discussion.—Examination of several specimens including
the holotype (G 40355, British Museum of Natural History)
from the Keele beds of the uppermost coal measures in
Northern Worcestershire (England), described by Cox (1926),
shows that the specimens from Karniowice are very similar in
their form and size to those from the type locality but differ in
being somewhat more slender.
Suborder ORTHURETRA Pilsbry, 1900
Family DENDROPUPIDAE Wenz, 1938
Genus DENDROPUPA Owen, 1859
DENDROPUPA ZARECZNYI Panow, 1936
Figure 4.8–4.10
Dendropupa zarecznyi Panow, 1936, p. 37, pl. 1, figs. 1–6.
Material examined.—One complete specimen (ISEZ-K/86),
five nearly complete and 16 poorly preserved fragments.
Dimensions (in mm).—Height: 9.75; Width 4.15; Number of
whorls ca. 6.5.
Description.—Shells very variable in shape: from tapering
cylindrical to pupiform, consisting of 6.5–7 whorls. Whorls
poorly convex, separated by a moderately impressed suture,
covered with very weak and somewhat irregular, radial striae
well visible on some fragments (Fig. 4.10). Sculpture of apical
whorls unknown because none of the specimens is sufficiently
well preserved. Aperture rounded with a broadly reflected
margin. The apertures of all the specimens are filled with
calcite sediment, difficult to remove, and details inside are not
visible, but there is no sign of any teeth.
Discussion.—Panow (1936) compared his D. zarecznyi with
D. vetusta (Dawson, 1855) and D. walchiarum Fischer, 1885 and
stated that the new species was more similar to the latter one,
described from the French Permian (Fischer, 1885). However,
according to Solem and Yochelson (1979), the systematic
distinction of D. walchiarum and D. vetusta is doubtful because
of their similar shape, size, sculpture and aperture. D. zarecznyi
differs from both these forms not only in size but also in having
fewer whorls, which are significantly higher.
Among the materials from Panow’s collection stored in the
Geological Museum in Cracow and labelled as D. zarecznyi, there
is neither the holotype nor paratypes. According to the museum’s
curator they were most probably lost during World War II.
Hereby we designate specimen ISEZ-K/86 as the neotype.
Family UNCERTAIN
Genus PROTODISCUS Solem and Yochelson, 1979
PROTODISCUS PRISCUS (Carpenter, 1867)
Figure 5.1–5.3
Zonites (Conulus) priscus Carpenter, 1867, in Dawson, 1867,
p. 331, figs. a–c, e.
Material examined.—One specimen partially embedded in a
chip of rock (ISEZ-K/59).
Dimensions (in mm).—Width 2.35; Height: shell 1.4, last
whorl 1.03.
 STWORZEWICZ ET AL.—PALEOZOIC GASTROPODS FROM POLAND 941

FIGURE 4—1–4, Anthracopupa ohioensis Whitfield, 1881: 1, the best preserved specimen ISEZ-K/30, 312; 2, specimen ISEZ-K/54 with body whorl
partly crushed, 312; 3, general shape of A. ohioensis according to the photograph in Solem and Yochelson (1979, pl. 1, fig. 19); 4, polished
section of rock piece from Karniowice with Anthracopupa shell, 310. 5–7, Anthracopupa britannica Cox, 1926; 5, specimen A-I-56/26 with
columellar tooth visible, 315; 6, specimen A-I-56/25 with the parietal tooth visible, 317; 7, three specimens of A. brittanica in one piece of rock
(ISEZ-K/1), showing the frequency of occurrence of the species, 34. 8–10, Dendropupa zarecznyi Panow, 1936: 8, neotype ISEZ-K/86 with the
upper whorls damaged, 38; 9, specimen ISEZ-K/60 with the last whorl and aperture partly damaged, 37; 10, the last two whorls of the neotype
ISEZ-K/86 showing detail of surface sculpture, 312.
942 JOURNAL OF PALEONTOLOGY, V. 83, NO. 6, 2009

FIGURE 5—1–3, Protodiscus priscus (Carpenter, 1867): 1, top view of specimen ISEZ-K/59, 318; 2, same specimen in side view, 324; 3, details of shell
sculpture, 328; 4, two halves of spherosiderite nodule with imprints of species A, ISEZ-K/93, 32; 5, shell reconstruction from a plastic mass mould
(species A), 32.5; 6, badly preserved shell imprint of species B, ISEZ-K/95, 34.

Description.—The specimen consists of 3 whorls, but it can Family UNCERTAIN

be inferred from the setting of the shell that a further part of
the last (?) whorl is partially embedded in the matrix together
with the whole umbilical part of the shell. Whorls well
rounded, separated by a deep suture. The shell is in large
part crystallized and the apical microsculpture unknown.
Postapical microsculpture visible in places even at low
magnification, especially when it is viewed at an adequate
angle and illumination (Fig. 5.2, 5.3). It consists of fine, rather
regular radial striae and widely spaced, finely marked ribs.
Discussion.—Solem and Yochelson (1979) referred Z.
priscus to the Discidae based on its shell sculpture, which
was, in their opinion, identical with the basic pattern found in
the extant Discidae. They argued that it could not be a
pupilloid snail because ‘‘Pupilloid taxa generally have apices
that are smooth or have microspiral grooves …’’ but some
pupilloids have apical sculpture of another type (Stworzewicz,
1999a). The sculpture of the teleoconch in the specimen from
Karniowice is similar to that of the type specimen, but the
protoconch is damaged and no trace of sculpture exists. The
last whorl of the type specimen increases rapidly in width, but
it is not visible in the specimen from Karniowice, perhaps
because it is partially embedded in hard rock. However, it is
also invisible in the specimen labeled as Archeozonites? priscus
from coal measures in Nova Scotia (Nov. 1881, G 100), stored
in the Natural History Museum, London, which was
compared with our material.
SPECIES A
Figure 5.4–5.5
Material examined.—Two specimens in the form of shell
imprints in siderite nodules (ISEZ-K/93, ISEZ-K/94).
Dimensions (in mm).—Height ca. 13; Width ca. 6
Description.—Shell regularly ovate in outline, consisting of
about five moderately convex whorls, separated by a scarcely
impressed suture. It seems that the apical whorls are broadly
rounded. Body whorl deflected downward before the aperture,
takes about two-thirds of the shell height. Aperture shape
difficult to reconstruct but it may be broadly rounded.
Discussion.—Because of the ovate shape, the specimens
resemble some living groups of terrestrial snails, both
prosobranch and pulmonate (for example some Poteriidae or
Pupinidae, but also Enidae). However, there are no characters
that would permit assessment of their affinity.
SPECIES B
Figure 5.6
Material examined.—One specimen in the form of shell
imprints in siderite nodule (ISEZ-K/95).
Dimensions (in mm).—Height ca. 15; Width ca. 7.5.
Description.—Specimen differs from the preceding form in
having more numerous (about seven to nine) but significantly
lower whorls and a more cylindrical shell. It seems that the
upper part of the spire is broadly conical, but because of being
 STWORZEWICZ ET AL.—PALEOZOIC GASTROPODS FROM POLAND
embedded in the siderite matrix it is hardly visible in front
view.
Discussion.—The imprint of this shell is badly preserved and
the plastic mould is also unsatisfying. In its general shape the
specimen bears a slight resemblance to some recent Cerion
species.
DISCUSSION
Recently, the systematic position of Paleozoic land gastro-
pods has been broadly discussed in attempts to resolve the
fundamental question of pulmonate evolution: which group
should have the basalmost position in the stylommatophoran
tree (Nordsieck, 1986; Tillier et al., 1995; Bandel, 2002;
Bouchet and Rocroi, 2005)? The problem mainly concerns the
two most numerous groups, the Anthracopupidae and
Dendropupidae. Solem and Yochelson (1979), in their
complete study of Upper Paleozoic land snails, regarded most
of them as terrestrial pulmonates. The same opinion had been
earlier expressed by Cox (1926 and literature cited therein),
albeit with a reservation that the condition of the described
material was poor, which made it difficult to conjecture about
phylogenetic relationships. Anthracopupa was also referred to
the Cyclophoridae or another prosobranch taxon (Knight et
al., 1960; Bandel, 2002), but such an affinity seems unlikely
because of the aperture structure (see Solem and Yochelson,
1979). Some recent authors express yet another and complete-
ly different opinion, namely that these Paleozoic snails have
very little in common with recent terrestrial gastropods, since
most probably no land snails survived the end-Permian mass
extinction (e.g., Wade et al., 2006). It should be remembered,
however, that not all animal taxa were equally affected by the
event. About 95% of marine species were exterminated during
the Permian extinction which, on the other hand, put an end to
‘‘only’’ 70% of terrestrial families (Goldberg et al., 2003; Lane,
2007). Furthermore, terrestrial invertebrates appear to have
survived the crisis better than marine animals or terrestrial
tetrapods; for example, only eight of the 27 orders of Paleozoic
insects became extinct, and nearly half of the remaining ones
survived till today (Labandeira and Sepkoski, 1993).
Solem and Yochelson (1979) classified Anthracopupa among
the Tornatellinidae and listed an array of arguments which, in
their opinion, were against placing Anthracopupa in the family
Ellobiidae—a view earlier suggested by Wenz (1938). However,
their arguments are not very convincing (see Nordsieck, 1986),
and subsequent authors most often placed Anthracopupa as a
distinct group within the Ellobioidea (for review of classification
systems, see Bouchet and Rocroi, 2005).
With respect to relationships (based on shell structure
similarities) with the Ellobiidae they were compared with
members of the genus Carychium, whereas actually the most
numerous characters are shared by Anthracopupa and
representatives of the genus Zospeum (cf. Bole, 1974), or—
which seems even more justified—the Tertiary genus Car-
ychiopsis. The genus was very abundantly represented during
the Paleocene [but not in the Paleozoic (!) as erroneously
suggested by Martins (1996) and Barker (2001)] and became
gradually extinct by the end of the Tertiary (Wenz, 1923;
Stworzewicz, 1999b). It is also likely that Carychium broti
Loriol, 1865 (Sandberger, 1870–1875), described from the
Upper Jurassic, was actually a member of Carychiopsis; this is
suggested by the copy of the figure reproduced in Sandberger
(1870–1875, pl. 1, fig. 33). The other possibility is relatedness
with the oldest known terrestrial ellobiids, Protocarychium
mirum Pan, 1982 and P. arcidentata Pan, 1982, described from
the Early Jurassic formation of China (Pan, 1982). On the
943
other hand, Anthracopupa was also referred to the Pupilli-
dae(?) (Yen, 1949).
Even assuming that Anthracopupa is an extinct ellobioid
taxon without descendants or an ancestor of a younger group,
there remains the question of the position of Ellobioidea in the
pulmonate phylogenetic tree. The problem has been recently
resurrected and repeatedly analysed based on both morpho-
logical and molecular characters, but relationships among
pulmonates remain unclear (Wade et al., 2000; Dayrat et al.,
2001; Dayrat and Tillier, 2002; Wade et al., 2006; Klussmann-
Kolb et al., 2008).
Possible relatives of the Dendropupidae were sought among
the Cyclophoroidea or Ellobioidea, but also among the Enidae
(Stylommatophora) (Solem and Yochelson, 1979; Batten,
1995) and Pupillidae(?) (Yen, 1949). However, Nordsieck’s
(1986) view that these snails constitute a distinct family within
the Orthurethra (Stylommatophora) seems to be the most
convincing; according to this author, the genus Protodiscus
described by Solem and Yochelson (1979) is also a member of
an orthurethran family. Solem and Yochelson (1979) placed
Protodiscus within the Discidae, but Nordsieck (1986) argued
that because of the shell form and microsculpture, P. priscus
was equally similar to the Pleurodiscidae. However, yet
another group should be considered; P. priscus is very similar
to the Valloniidae in its shell form and especially in its surface
sculpture. Many members of Vallonia and also Planogyra have
a shell surface covered by closely spaced, delicate striae and
more widely spaced, more pronounced riblets. The oldest
Valloniidae are known only from as late as the Paleocene
(Gerber, 1996), but there is no doubt the group must have
arisen much earlier, as indicated by a typically valloniid shell
of Vallonia sparnacensis (Deshayes, 1863) found in France
(Deshayes, 1863).
CONCLUSIONS
Study of Upper Carboniferous and Lower Permian con-
tinental sediments of the Silesian-Cracow area has yielded two
taxa from the Late Carboniferous and four species of Early
Permian terrestrial snails.
The described snail assemblages provide new data on the
distribution of Late Paleozoic terrestrial gastropods and their
evolutionary context. It has become obvious that the
paleogeographical range of Anthracopupa and Protodiscus
extended from North America to the European part of the
Pangean supercontinent and was considerably broader than
had been previously assumed. It means that taxonomic
diversity of the oldest known land snails from this part of
Pangea was not any less than those in North America.
Furthermore, the time range of Anthracopupa and Protodiscus
must be shifted up to the Early Permian.
The accompanying other organisms (plants, insects, crusta-
cean, and bivalves, both freshwater and terrestrial) together
with sedimentological indicators, make possible the relatively
precise reconstruction of the habitats of the first terrestrial
gastropods. According to these data, the swamp facies harbored
a rich malacofauna that inhabitated the lowland peat area.
ACKNOWLEDGMENTS
We thank B. Kietlińska-Michalik (The Geological Museum,
PAS, Cracow) for the loan of Panow’s specimens and J. Todd
(The Natural History Museum, London) for the loan of the
British material of Paleozoic snails. The snail-bearing sphero-
siderites were kindly made available by D. Wojciechowski.
Professors A. Riedel and R. A. D. Cameron helped us during
field work. We thank also G. Kaim and B. Kołodziej for their
944 JOURNAL OF PALEONTOLOGY, V. 83, NO. 6, 2009
photographic assistance. We are particulary grateful to A.
Nutzel and D. Rohr for their helpful reviews.
The studies were financed by the grant MNiI no. 2P04C 011
27.
REFERENCES
BANDEL, K. 2002. Reevaluation and classification of Carboniferous and
Permian Gastropoda belonging to the Caenogastropoda and their
relation. Mitteilugen aus dem Geologisch-Paläontologisches Institut der
Universität Hamburg, 86:81–188.
BARKER, G. M. 2001. Gastropods on land: phylogeny, diversity and
adaptative morphology, p. 1–146. In G. M. Barker (ed.), The Biology of
Terrestrial Molluscs. CAB International, Oxford.
BATTEN, R. L. 1995. Pennsylvanian (Morrowan) Gastropods from the
Magdalena Formation of the Hueco Mountains, Texas. American
Museum Novitates, 3122:1–46.
BOLE, J. 1974. Rod Zospeum Bourguignat 1856 (Gastropoda, Ellobiidae)
v Jugoslaviji. Razprave, Slovenska Akademija Znanosti in Umetnosti,
17:251–291.
BOUCHET, P. AND J-P. ROCROI. 2005. Classification and Nomenclator of
Gastropod Families. Malacologia, 47:1–397.
COX, L. R. 1926. Anthracopupa brittanica sp. nov., a Land Gastropod
from the Red Beds of the Uppermost Coal-Measures of Northern
Worcestershire. Quarterly Journal of the Geological Society, 82:401–
410.
CUVIER, G. L. C. 1797. Tableau Èlèmentaire de l’Histoire Naturelle des
Animaux [des Mollusques]. Baudonin, Paris, 710 p.
ĆWIŻEWICZ, M. AND J. SZULC. 1989. Warunki klimatyczne środowiska
sedymentacji martwicy karniowickiej. Przegla˛d Geologiczny, 37:180–
187.
DAWSON, J. W. 1855. Acadian geology; an account of the geological
structure and mineral resources of Nova Scotia and portions of the
neighboring provinces of British America. Oliver and Boyd, Edinburgh,
388 p.
DAWSON, J. W. 1867. On the discovery of a new pulmonate mollusk
(Zonites (Conulus) priscus, Cpr.) in the coal-formation of Nova Scotia,
with a description of the species by Philip P. Carpenter. Quarterly
Journal of the Geological Society of London, 23:330–333.
DAYRAT, B. AND S. TILLIER. 2002. Evolutionary relationships of
euthyneuran gastropods (Mollusca): a cladistic re-evaluation of
morphological characters. Zoological Journal of the Linnean Society,
135:403–470.
DAYRAT, B., A. TILLIER, G. LECOINTRE, AND S. TILLIER. 2001. New
Clades of Euthyneuran Gastropods (Mollusca) from 28S rRNA
Sequences. Molecular Phylogenetics and Evolution, 19:225–235.
DESHAYES, G. P. (1861–1864). Description des Animaux sans vertèbres du
Bassin de Paris, 2. Paris.
FALCON-LANG, H. J., M. J. BENTON, S. J. BRADDY, AND S. J. DAVIES.
2006. The Pennsylvanian tropical biome reconstructed from the Joggins
Formation of Nova Scotia, Canada. Journal of the Geological Society,
London, 163:561–576.
FILIPIAK, P. AND P. KRAWCZYŃSKI. 1996. Westphalian xiphosurans
(Chelicerata) from the Upper Silesia Coal Basin of Sosnowiec, Poland.
Acta Palaeontologica Polonica, 41:413–425.
FISCHER, P. 1885. Description d’une nouvelle espèce de Dendropupa, du
terrain Permien de Saône-et-Loire. Journal de Conchyliologie, 33:99–
105.
GERBER, J. 1996. Revision der Gattung Vallonia Risso 1826 (Mollusca:
Gastropoda: Valloniidae). Schriften zur Malakozoologie aus dem Haus
der Natur – Cismar, 8:1–227.
GOLDBERG, S., J. MA, AND J. O’DONOHUE. 2003. Mass Extinction. The
Traprock, 2:5–18.
HASZPRUNAR, G. AND G. HUBER. 1990. On the central nervous system of
Smeagolidae and Rhodopidae, two families questionably allied with the
Gymnomorpha (Gastropoda: Euthyneura). Journal of Zoology,
220:185–199.
HEBERT, B. L. AND J. H. CALDER. 2004. On the discovery of a unique
terrestrial faunal assemblage in the classic Pennsylvanian section at
Joggins, Nova Scotia. Canadian Journal of Earth Science, 41:247–254.
KLUSSMANN-KOLB, A., A. DINAPOLI, K. KUHN, B. STREIT, AND C.
ALBRECHT. 2008. From sea to land and beyond – New insights into the
evolution of euthyneuran Gastropoda (Mollusca). BMC Evolutionary
Biology, 8:57.
KNIGHT, J. B., R. L. BATTEN, E. L. YOCHELSON, AND L. R. COX. 1960.
Paleozoic and some Mesozoic Caenogastropoda and Opistobranchia, p.
1310–1324. In R.C. Moore (ed.) Treatise on Invertebrate Paleontology,
Part I, Mollusca 1. Geological Society of America, Inc. and University
of Kansas Press, Lawrence.
KRAWCZYŃSKI, P., P. FILIPIAK, AND M. GWOŹDZIEWICZ. 1997. Zespół
skamieniałości z karbońskich sferosyderytów (westfal A) NE cze˛ści
Górnośla˛skiego Zagłe˛bia We˛glowego. Przegla˛d Geologiczny, 45:1271–
1274.
LABANDEIRA, C. C. AND J. J. SEPKOSKI JR. 1993. Insect Diversity in the
Fossil Record. Science, 261:310–315.
LANE, N. 2007. Reading the book of death. Nature, 448:122–125.
LINNAEUS, C. 1758. Systema Naturae per Regna Tria Naturae, Editio
decima, reformata, 1, Regnum Animale. Laurentii Salvii, Stockholm,
824 p.
LIPIARSKI, I. 1971. Dolnopermska flora martwicy karniowickiej koło
Krakowa. Instytut Geologiczny, Prace, 58:5–80.
LORIOL, P. de AND A. JACCARD. 1865. Etude géologique et paléontolo-
gique de la formation d’eau douce infracrétacée du Jura, et en
particulier de Villers-le-Lac. Mémoires de la Société de Physique et
d’Histoire naturelle de Genève, 18:1–68.
MARTINS, A. M. F. 1996. Anatomy and systematics of the Western Atlantic
Ellobiidae (Gastropoda: Pulmonata). Malacologia, 37:163–332.
NORDSIECK, H. 1986. The system of the Stylommatophora (Gastropoda),
with special regard to the systematic position of the Clausiliidae, II.
Importance of the shell and distribution. Archiv für Molluskenkunde,
117:93–116.
OPLUŠTIL, S. 2004. Late Carboniferous tectono-sedimentary evolution
and related terrestrial biotic changes on the North Variscan and
Appalachian forelands, and adjacent paralic and continental basins.
Geologica Balcanica, 34:51–67.
PACYNA, G. 2003. Carboniferous fructifications from a new locality in
Sosnowiec – Preliminary report. Proceedings XXVI Symposium
Geology of Coal-bearing strata of Poland, Cracow, 121–125.
PACYNA, G. AND D. ZDEBSKA. 2001. Upper Carboniferous plant
macrofossils from sideritic concretions of the Upper Silesian Coal
Basin of Sosnowiec. Preliminary report. Materiały XXIV Sympozjum
Geologia formacji we˛glonośnych Polski, Kraków, 75–79.
PACYNA, G. AND D. ZDEBSKA. 2002. Upper Carboniferous plant
macrofossils from sideritic concretions in Sosnowiec (Upper Silesian
Coal Basin) and Mazon Creek (Illinois, USA). Proceedings, XXV
Symposium Geology of Coal-bearing strata of Poland, Cracow, 123–127.
PAN, H. 1982. Late Triassic-Early Jurassic Gastropods from Eastern
Hunan and Northeastern Guangxi. Memoirs of Nanjing Institute of
Geology and Paleontology, 17:85–112.
PANOW, E. 1936. Permokarbońska fauna martwicy karniowickiej.
Rocznik Polskiego Towarzystwa Geologicznego, 12:36–41.
PFEIFFER, L. 1854. Synopsis Auriculaceorum. Malakozoolologische
Blätter, 1:145–156.
PILSBRY, H. A. 1900. On the zoological position of Partula and
Achatinella. Proceedings of the Academy of Natural Sciences of
Philadelphia, 52:561–567.
RACIBORSKI, M. 1891. Permokarbońska flora karniowickiego wapienia.
Rozprawy Akademii Umieje˛tności, Wydział matematyczno-przyrod-
niczy, Ser. 2, 1:353–394.
SANDBERGER, F. (1870–1875). Die Land- und Süsswasser-conchylien der
Vorwelt. C. W. Kreidel’s Verlag, Wiesbaden. 1000 p.
SOLEM, A. AND E. L. YOCHELSON. 1979. North American Paleozoic Land
Snails, With a Summary of Other Paleozoic Nonmarine Snails. U.S.
Geological Survey Professional Paper, 1072:1–42.
STWORZEWICZ, E. 1999a. Miocene land snails from Bełchatów (Central
Poland), IV: Pupilloidea (Gastropoda Pulmonata). Systematic, bio-
stratigraphic and paleoecological studies. Folia Malacologica, 7:133–
170.
STWORZEWICZ, E. 1999b. Miocene land snails from Bełchatów (Central
Poland), III. Carychiinae (Gastropoda; Pulmonata: Ellobiidae). Pa-
läontologische Zeitschrift, 73:261–276.
SZULC, J. AND M. ĆWIŻEWICZ. 1989. The Lower Permian freshwater
carbonates of the Slawków Graben, Southern Poland: sedimentary
facies context and stable isotope study. Paleogeography, Paleoclima-
tology, Paleoecology, 70:107–120.
TILLIER, S., M. MASSELOT, AND A. TILLIER. 1995 [1996]. Phylogenetic
relationships of the pulmonate gastropods from rRNA sequences, and
tempo and age of the stylommatophoran radiation, p. 267–284. In J. D.
Taylor (ed.) Origin and evolutionary radiation of the Mollusca. Oxford
University Press.
WADE, C. M., P. B. MORDAN, AND B. CLARKE. 2000. A phylogeny of the
land snails (Gastropoda: Pulmonata). Proceedings of the Royal Society
of London, 268:413–422.
WADE, C. M., P. B. MORDAN, AND F. NAGGS. 2006. Evolutionary
relationships among the Pulmonate land snails and slugs (Pulmonata,
Stylommatophora). Biological Journal of the Linnean Society, 87:593–
610.
WENZ, W. 1923. Gastropoda extramarina tertiaria, p. 1069–1420. In C.
Diener (ed.) Fossilium Catalogus, I: Animalia, 21. W. Junk, Berlin.
 STWORZEWICZ ET AL.—PALEOZOIC GASTROPODS FROM POLAND 945

WENZ, W. 1938. Gastropoda I: Allgemeiner Teil und Prosobranchia, p. 1–
480. In O. H. Schindewolf (ed.) Handbuch der Paläozoologie, Band 6.
Borntraeger, Berlin.
WHITFIELD, R. P. 1881. Notice of a new genus and species of air-
breathing mollusk from the Coal-Measures of Ohio, and observations
on Dawsonella. American Journal of Sciences, ser. 3, 21, 125–
128.
YEN, T. C. 1949. Review of Paleozoic non-marine Gastropods and a
description of a new genus from the Carboniferous rocks of Scotland.
Journal of Molluscan Studies, 27:235–240.
ZARE˛CZNY, S. 1894. Mapa geologiczna okolic Krakowa i Chrzanowa.
Kraków, 280 p.
ACCEPTED 12 JULY 2009