You are on page 1of 8

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- spherosiderites. Out of several thousand siderite nodules
T HE ADVENT
tiation in the earliest stages of their evolution have long
been of great interest to malacologists, this being among the
collected from spoil-heaps in Sosnowiec, only a few dozen
contained well-preserved fragments of plants and animals
crucial problems of gastropod evolutionary relationships. The (Krawczyński et al., 1997). Molluscs were found in only four
extreme paucity of the fossil record does not enable a reliable concretions, one of which contained bivalve remains, while the
reconstruction of the early history of the terrestrial malaco- remaining ones held three snail imprints (ISEZ-K/93–ISEZ-K/
fauna. The Upper Silesian–Cracow area is among the few 95) described here for the first time.
regions with a fossil record of land snails from the Late Because of the great diversity of spherosiderite fossils, the
Palezoic. The most abundant data come from the Upper site in Sosnowiec is very valuable, especially from a faunistic
Palezoic rocks of the United States and Canada. The existing viewpoint, ranking beside the two most famous sites with
fossil record indicates that at the end of the Paleozoic, remains of Carboniferous plants and animals in Mazon Creek
terrestrial snails were already quite diverse. Solem and (Illinois, USA) and at Joggins (Nova Scotia, Canada) (Pacyna
Yochelson (1979) assigned them to three out of the six orders and Zdebska, 2002; Hebert and Calder, 2004; Falcon-Land et
known today and recognised 10 pulmonate and one terrestrial al., 2006). Faunal assemblages known from other European
prosobranch species, whereas Bandel (2002) included the Late coal deposits are distinctly poorer.
Paleozoic gastropods in a new order, the Procyclophorida. The snails found in three spherosiderite concretions are shell
Based on the previous studies, the total number of terrestrial imprints, and the shell form was reconstructed as moulds
Paleozoic snails worldwide does not exceed 25. using plastic mass. In only one case was the condition of the
To date, the only Polish contribution to these studies was a imprint sufficiently good to permit a reconstruction of the
description of Dendropupa zarecznyi by Panow (1936) from exact shape of the shell and its aperture.
the Lower Permian freshwater limestones of Karniowice, near Lower Permian freshwater carbonates – ‘‘Karniowice Trav-
Cracow. The present study yields many new specimens, ertine’’ (Fig. 1, locality 2).—In the region of Cracow, Zare˛czny
representing some additional species and allowing a re- (1894) found Paleozoic deposits containing gastropod shells as
evaluation of the previous results of study of the Late well as numerous plant remains. These materials originating
Paleozoic continental gastropods. from the so-called ‘‘Karniowice travertine’’ served as the basis
Beside the previously known Early Permian malacofauna, for a paper by Panow (1936), containing a description of a new
there was no published information on gastropod remains from species, while mention of some other snails from the site
other sites from southern Poland. The recent findings of the appeared in Solem and Yochelson (1979).
gastropods from the Upper Carboniferous terrestrial sediments Carbonate deposits in the region of Krzeszowice near
(Filipiak and Krawczyński, 1996; Krawczyński et al., 1997) Cracow (50u099N, 19u339E), often collectively referred to as
provided specimens representing probably two snail species. ‘‘Karniowice travertine,’’ are especially interesting among
other Lower Permian sedimentary rocks of the Cracow region.
LOCATION AND MATERIAL Recent studies (Szulc and Ćwiz_ewicz, 1989; Ćwiz_ewicz and
Carboniferous siderite nodules from Sosnowiec (Fig. 1, Szulc, 1989) showed that the spectrum of carbonate rocks is
locality 1).—The snail-bearing spherosiderite concretions were much broader than travertine formations sensu stricto; other
found on a spoil-heap of the Pora˛bka–Klimontów coal mine rocks include calcareous deposits formed in small stagnant
in Sosnowiec, in the NE. part of the Upper Silesian Mining water bodies (semi-limnic), in streams feeding these ponds,
Region (50u179N, 19u079E). Their age was estimated as and calcareous deposits of swamp facies with numerous
Westfalian A by means of palynological and macrofloral calcified remains of vascular plants (Fig. 2). Based on a very
analysis (Krawczyński et al., 1997; Pacyna, 2003). The rich assemblage of such plants, Lipiarski (1971) described their
concretion-bearing rocks also comprise other freshwater and habitat as a topogenous fen. The vascular plant assemblage
terrestrial fauna, such as crustaceans, bivalves, insects, and includes both Carboniferous and Permian components, which
remains of terrestrial plants of the genera Calamites, Sigillaria, makes it difficult to unambiguosuly determine the age of the
and Lepidostrobus. Karniowice travertine (Raciborski, 1891). The ultimate
938
STWORZEWICZ ET AL.—PALEOZOIC GASTROPODS FROM POLAND 939

FIGURE 2—Scheme of the Lower Permian carbonate facies distribution


FIGURE 1—Location map of the gastropod occurrences. 1, Upper of the Karniowice Travertine. 1, Lower Carboniferous basement rocks; 2,
Carboniferous gastropods; 2, Lower Permian gastropods. fanglomerates with pyroclastics; 3, spring deposits –travertine sensu
stricto; 4, paleosoils with caliche crusts.
criterion for classifying the travertine as Lower Permian was
the high proportion of new components that were absent from the Joggins Formation deposits, were characterized by a diverse
Carboniferous assemblages (Lipiarski, 1971). Survival of vegetation comprising pteridosperms, ferns, lycopsids, cordai-
Carboniferous tropical flora in the generally semi-arid climate taleans and calamiteans (Falcon-Land et al., 2006). One facies
of the early Permian was possible only due to the presence of a of these deposits comprises siderite nodules with tetrapod
vast system of karstic springs that fed an extensive oasis. skeletal material and some other animals. Siderite nodules from
As mentioned above, the first Paleozoic land snails were Sosnowiec also contain a rich faunal assemblage, including
found over a hundred years ago by Zare˛czny (1894). Based on crustaceans and molluscs. The plant macrofossils represent
his materials, Panow (1936) described Dendropupa zarecznyi. typical Carboniferous plant taxa, such as lycopods, sphenop-
He mentioned also several other specimens of other species sids, pteridophylls, cordaites and conifers (Pacyna and
collected by him, but these have never been examined or Zdebska, 2001, 2002; Pacyna, 2003). The animal and plant
described. General information on Dendropupa snails from fossils represent two environments—terrestrial and aquatic
Karniowice was provided by Solem and Yochelson (1979). (Filipiak and Krawczyński, 1996; Krawczyński et al., 1997).
Yochelson visited the site in 1968 and collected a few more The plant remains, insects and snails indicate terrestrial
specimens, stored in the National Museum of Natural History habitats; the crustaceans are freshwater forms, while bivalves
of Smithsonian Institution in Washington. inhabit both fresh and brackish waters.
The material for this study was collected by the authors The calcified floral remnants in the freshwater limestones
from 2004 to 2007. It includes 92 limestone samples with snail from Karniowice include many taxa also recorded from
fossils (ISEZ-K/1–ISEZ-K/92), from one to seven per frag- deposits of the Joggins Formation. These plant assemblages
ment (a total of 170 specimens). Unfortunately, poor represent mainly a hygrophilous or nearly-hygrophilous
preservation of the specimens often precludes precise species vegetation type characterictic of Carboniferous and early
identification. Only a few specimens show traces of shell Permian peat associations (Lipiarski, 1971). Based on their
surface sculpture. The collection is kept in the Institute of stable isotope study, Szulc and Ćwiz_ewicz (1989) pointed out
Systematics and Evolution of Animals, Polish Academy of that the Lower Permian Karniowice freshwater carbonates
Sciences, Cracow. Additional material comes from Panow’s were deposited in subtropical conditions comparable to those
collection stored in the Geological Museum PAS, Cracow of the present-day subtropics. These carbonates are composed
(A-I-56/1–A-I-56/27). of travertines, lacustrine, and palustrine limestones deposited
All measurements were taken with Nikon measurescope in a spring-fed oasis basin. Such conditions were favorable for
MM-11, accuracy 0.01 mm. a range of animals, including the earliest terrestrial gastro-
pods, which required a wetland environment at the early
PALEOGEOGRAPHICAL AND PALEOENVIRONMENTAL CONTEXT stages of their terrestrial life. It is also noteworthy that the
At the end of the Carboniferous, a collision of continental snails discussed occur only in the palustrine facies of the
plates led to the formation of the large supercontinent Pangea Karniowice Travertine limetsones.
and to origin of the Variscian orogenic belt, with a vast foreland The high proportion of small snails is a characteristic
basin system filled by coal-bearing molasse deposits (Fig. 3). feature of the Karniowice assemblage, while the considerably
The continental plate repositioning resulted in changes to larger D. zarecznyi is represented by only few specimens. It
atmospheric circulation and moisture distribution and brought is interesting that D. zarecznyi was always found together
about floristic changes (Opluštil, 2004). The paleogeographical with large plant fragments. In contrast, small shells were
position of the Variscan belt imposed tropical climatic accompanied by fine plant fragments. These facts may indicate
conditions in the late Carboniferous, generally hot and humid. either different microhabitat requirements or slightly different
Wetland terrestrial ecosystems, reconstructed on the basis of taphonomic modes.
940 JOURNAL OF PALEONTOLOGY, V. 83, NO. 6, 2009

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
FIGURE 3—Paleogeographic position of the discussed areas. 1, Cracow-
Upper Silesian area; 2, Joggins Formation of Nova Scotia. Dendropupa zarecznyi Panow, 1936, p. 37, pl. 1, figs. 1–6.
Material examined.—One complete specimen (ISEZ-K/86),
SYSTEMATIC PALEONTOLOGY five nearly complete and 16 poorly preserved fragments.
Dimensions (in mm).—Height: 9.75; Width 4.15; Number of
Phylum MOLLUSCA Linnaeus, 1758
whorls ca. 6.5.
Class GASTROPODA Cuvier, 1797 Description.—Shells very variable in shape: from tapering
Order EUPULMONATA Haszprunar and Huber, 1990 cylindrical to pupiform, consisting of 6.5–7 whorls. Whorls
Superfamily ELLOBIOIDEA L. Pfeiffer, 1854 poorly convex, separated by a moderately impressed suture,
Family ANTHRACOPUPIDAE Wenz, 1938 covered with very weak and somewhat irregular, radial striae
Genus ANTHRACOPUPA Whitfield, 1881 well visible on some fragments (Fig. 4.10). Sculpture of apical
ANTHRACOPUPA OHIOENSIS Whitfield, 1881 whorls unknown because none of the specimens is sufficiently
Figure 4.1–4.4 well preserved. Aperture rounded with a broadly reflected
Anthracopupa ohioensis Whitfield, 1881, p. 126, figs. 1–4. margin. The apertures of all the specimens are filled with
calcite sediment, difficult to remove, and details inside are not
Material examined.—Three specimens ISEZ-K/11, ISEZ-K/
visible, but there is no sign of any teeth.
54, ISEZ-K/30 (only the latter in relatively good condition). Discussion.—Panow (1936) compared his D. zarecznyi with
Dimensions (in mm).—Height: 4.40, height of last whorl: D. vetusta (Dawson, 1855) and D. walchiarum Fischer, 1885 and
3.15; Width: 3.03; Number of whorls ca. 4.5. stated that the new species was more similar to the latter one,
Description.—Best-preserved shell of ovoid shape, consist- described from the French Permian (Fischer, 1885). However,
ing of 4.5 moderately convex whorls separated by rather according to Solem and Yochelson (1979), the systematic
shallow suture. Rounded aperture filled with calcite sediment, distinction of D. walchiarum and D. vetusta is doubtful because
difficult to remove, hence no apertural teeth visible. In another of their similar shape, size, sculpture and aperture. D. zarecznyi
specimen, due to the body whorl being partly crushed, differs from both these forms not only in size but also in having
something like a columellar tooth can be seen inside fewer whorls, which are significantly higher.
(Fig. 4.2). Similar structure also visible in the polished section Among the materials from Panow’s collection stored in the
of rock piece with Anthracopupa shell (Fig. 4.4). Aperture Geological Museum in Cracow and labelled as D. zarecznyi, there
margin broken so that the outer lip thickening is difficult to is neither the holotype nor paratypes. According to the museum’s
define. The surface sculpture also invisible. curator they were most probably lost during World War II.
Discussion.—The specimens from Karniowice are strikingly Hereby we designate specimen ISEZ-K/86 as the neotype.
similar to the shorter and more tumid shells of A. ohioensis,
presented by Solem and Yochelson (1979, pl. 1, fig. 19), in Family UNCERTAIN
having the same shell shape and form of the aperture Genus PROTODISCUS Solem and Yochelson, 1979
(Fig. 4.3); however the latter are generally somewhat smaller. PROTODISCUS PRISCUS (Carpenter, 1867)
Figure 5.1–5.3
ANTHRACOPUPA BRITANNICA Cox, 1926
Figure 4.5–4.7 Zonites (Conulus) priscus Carpenter, 1867, in Dawson, 1867,
p. 331, figs. a–c, e.
Anthracopupa britannica Cox, 1926, p. 407, figs. 1a, 1b, 2a, 2b. Material examined.—One specimen partially embedded in a
Material examined.—five specimens in relatively good chip of rock (ISEZ-K/59).
condition (ISEZ-K/71, ISEZ-K/55, ISEZ-K/1a–c) and 115 Dimensions (in mm).—Width 2.35; Height: shell 1.4, last
fragments of shells. 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 SPECIES A
the last (?) whorl is partially embedded in the matrix together Figure 5.4–5.5
with the whole umbilical part of the shell. Whorls well Material examined.—Two specimens in the form of shell
rounded, separated by a deep suture. The shell is in large imprints in siderite nodules (ISEZ-K/93, ISEZ-K/94).
part crystallized and the apical microsculpture unknown. Dimensions (in mm).—Height ca. 13; Width ca. 6
Postapical microsculpture visible in places even at low Description.—Shell regularly ovate in outline, consisting of
magnification, especially when it is viewed at an adequate about five moderately convex whorls, separated by a scarcely
angle and illumination (Fig. 5.2, 5.3). It consists of fine, rather impressed suture. It seems that the apical whorls are broadly
regular radial striae and widely spaced, finely marked ribs. rounded. Body whorl deflected downward before the aperture,
Discussion.—Solem and Yochelson (1979) referred Z. takes about two-thirds of the shell height. Aperture shape
priscus to the Discidae based on its shell sculpture, which difficult to reconstruct but it may be broadly rounded.
was, in their opinion, identical with the basic pattern found in Discussion.—Because of the ovate shape, the specimens
the extant Discidae. They argued that it could not be a resemble some living groups of terrestrial snails, both
pupilloid snail because ‘‘Pupilloid taxa generally have apices prosobranch and pulmonate (for example some Poteriidae or
that are smooth or have microspiral grooves …’’ but some Pupinidae, but also Enidae). However, there are no characters
pupilloids have apical sculpture of another type (Stworzewicz, that would permit assessment of their affinity.
1999a). The sculpture of the teleoconch in the specimen from
Karniowice is similar to that of the type specimen, but the SPECIES B
protoconch is damaged and no trace of sculpture exists. The Figure 5.6
last whorl of the type specimen increases rapidly in width, but Material examined.—One specimen in the form of shell
it is not visible in the specimen from Karniowice, perhaps imprints in siderite nodule (ISEZ-K/95).
because it is partially embedded in hard rock. However, it is Dimensions (in mm).—Height ca. 15; Width ca. 7.5.
also invisible in the specimen labeled as Archeozonites? priscus Description.—Specimen differs from the preceding form in
from coal measures in Nova Scotia (Nov. 1881, G 100), stored having more numerous (about seven to nine) but significantly
in the Natural History Museum, London, which was lower whorls and a more cylindrical shell. It seems that the
compared with our material. upper part of the spire is broadly conical, but because of being
STWORZEWICZ ET AL.—PALEOZOIC GASTROPODS FROM POLAND 943

embedded in the siderite matrix it is hardly visible in front other hand, Anthracopupa was also referred to the Pupilli-
view. dae(?) (Yen, 1949).
Discussion.—The imprint of this shell is badly preserved and Even assuming that Anthracopupa is an extinct ellobioid
the plastic mould is also unsatisfying. In its general shape the taxon without descendants or an ancestor of a younger group,
specimen bears a slight resemblance to some recent Cerion there remains the question of the position of Ellobioidea in the
species. pulmonate phylogenetic tree. The problem has been recently
resurrected and repeatedly analysed based on both morpho-
DISCUSSION
logical and molecular characters, but relationships among
Recently, the systematic position of Paleozoic land gastro- pulmonates remain unclear (Wade et al., 2000; Dayrat et al.,
pods has been broadly discussed in attempts to resolve the 2001; Dayrat and Tillier, 2002; Wade et al., 2006; Klussmann-
fundamental question of pulmonate evolution: which group Kolb et al., 2008).
should have the basalmost position in the stylommatophoran Possible relatives of the Dendropupidae were sought among
tree (Nordsieck, 1986; Tillier et al., 1995; Bandel, 2002; the Cyclophoroidea or Ellobioidea, but also among the Enidae
Bouchet and Rocroi, 2005)? The problem mainly concerns the (Stylommatophora) (Solem and Yochelson, 1979; Batten,
two most numerous groups, the Anthracopupidae and 1995) and Pupillidae(?) (Yen, 1949). However, Nordsieck’s
Dendropupidae. Solem and Yochelson (1979), in their (1986) view that these snails constitute a distinct family within
complete study of Upper Paleozoic land snails, regarded most the Orthurethra (Stylommatophora) seems to be the most
of them as terrestrial pulmonates. The same opinion had been convincing; according to this author, the genus Protodiscus
earlier expressed by Cox (1926 and literature cited therein), described by Solem and Yochelson (1979) is also a member of
albeit with a reservation that the condition of the described an orthurethran family. Solem and Yochelson (1979) placed
material was poor, which made it difficult to conjecture about Protodiscus within the Discidae, but Nordsieck (1986) argued
phylogenetic relationships. Anthracopupa was also referred to that because of the shell form and microsculpture, P. priscus
the Cyclophoridae or another prosobranch taxon (Knight et was equally similar to the Pleurodiscidae. However, yet
al., 1960; Bandel, 2002), but such an affinity seems unlikely another group should be considered; P. priscus is very similar
because of the aperture structure (see Solem and Yochelson, to the Valloniidae in its shell form and especially in its surface
1979). Some recent authors express yet another and complete- sculpture. Many members of Vallonia and also Planogyra have
ly different opinion, namely that these Paleozoic snails have a shell surface covered by closely spaced, delicate striae and
very little in common with recent terrestrial gastropods, since more widely spaced, more pronounced riblets. The oldest
most probably no land snails survived the end-Permian mass Valloniidae are known only from as late as the Paleocene
extinction (e.g., Wade et al., 2006). It should be remembered, (Gerber, 1996), but there is no doubt the group must have
however, that not all animal taxa were equally affected by the arisen much earlier, as indicated by a typically valloniid shell
event. About 95% of marine species were exterminated during of Vallonia sparnacensis (Deshayes, 1863) found in France
the Permian extinction which, on the other hand, put an end to (Deshayes, 1863).
‘‘only’’ 70% of terrestrial families (Goldberg et al., 2003; Lane,
2007). Furthermore, terrestrial invertebrates appear to have CONCLUSIONS
survived the crisis better than marine animals or terrestrial Study of Upper Carboniferous and Lower Permian con-
tetrapods; for example, only eight of the 27 orders of Paleozoic tinental sediments of the Silesian-Cracow area has yielded two
insects became extinct, and nearly half of the remaining ones taxa from the Late Carboniferous and four species of Early
survived till today (Labandeira and Sepkoski, 1993). Permian terrestrial snails.
Solem and Yochelson (1979) classified Anthracopupa among The described snail assemblages provide new data on the
the Tornatellinidae and listed an array of arguments which, in distribution of Late Paleozoic terrestrial gastropods and their
their opinion, were against placing Anthracopupa in the family evolutionary context. It has become obvious that the
Ellobiidae—a view earlier suggested by Wenz (1938). However, paleogeographical range of Anthracopupa and Protodiscus
their arguments are not very convincing (see Nordsieck, 1986), extended from North America to the European part of the
and subsequent authors most often placed Anthracopupa as a Pangean supercontinent and was considerably broader than
distinct group within the Ellobioidea (for review of classification had been previously assumed. It means that taxonomic
systems, see Bouchet and Rocroi, 2005). diversity of the oldest known land snails from this part of
With respect to relationships (based on shell structure Pangea was not any less than those in North America.
similarities) with the Ellobiidae they were compared with Furthermore, the time range of Anthracopupa and Protodiscus
members of the genus Carychium, whereas actually the most must be shifted up to the Early Permian.
numerous characters are shared by Anthracopupa and The accompanying other organisms (plants, insects, crusta-
representatives of the genus Zospeum (cf. Bole, 1974), or— cean, and bivalves, both freshwater and terrestrial) together
which seems even more justified—the Tertiary genus Car- with sedimentological indicators, make possible the relatively
ychiopsis. The genus was very abundantly represented during precise reconstruction of the habitats of the first terrestrial
the Paleocene [but not in the Paleozoic (!) as erroneously gastropods. According to these data, the swamp facies harbored
suggested by Martins (1996) and Barker (2001)] and became a rich malacofauna that inhabitated the lowland peat area.
gradually extinct by the end of the Tertiary (Wenz, 1923;
Stworzewicz, 1999b). It is also likely that Carychium broti ACKNOWLEDGMENTS
Loriol, 1865 (Sandberger, 1870–1875), described from the We thank B. Kietlińska-Michalik (The Geological Museum,
Upper Jurassic, was actually a member of Carychiopsis; this is PAS, Cracow) for the loan of Panow’s specimens and J. Todd
suggested by the copy of the figure reproduced in Sandberger (The Natural History Museum, London) for the loan of the
(1870–1875, pl. 1, fig. 33). The other possibility is relatedness British material of Paleozoic snails. The snail-bearing sphero-
with the oldest known terrestrial ellobiids, Protocarychium siderites were kindly made available by D. Wojciechowski.
mirum Pan, 1982 and P. arcidentata Pan, 1982, described from Professors A. Riedel and R. A. D. Cameron helped us during
the Early Jurassic formation of China (Pan, 1982). On the 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. KRAWCZYŃSKI, P., P. FILIPIAK, AND M. GWOŹDZIEWICZ. 1997. Zespół
skamieniałości z karbońskich sferosyderytów (westfal A) NE cze˛ści
Nutzel and D. Rohr for their helpful reviews. Górnośla˛skiego Zagłe˛bia We˛glowego. Przegla˛d Geologiczny, 45:1271–
The studies were financed by the grant MNiI no. 2P04C 011 1274.
27. LABANDEIRA, C. C. AND J. J. SEPKOSKI JR. 1993. Insect Diversity in the
Fossil Record. Science, 261:310–315.
REFERENCES LANE, N. 2007. Reading the book of death. Nature, 448:122–125.
BANDEL, K. 2002. Reevaluation and classification of Carboniferous and LINNAEUS, C. 1758. Systema Naturae per Regna Tria Naturae, Editio
Permian Gastropoda belonging to the Caenogastropoda and their decima, reformata, 1, Regnum Animale. Laurentii Salvii, Stockholm,
relation. Mitteilugen aus dem Geologisch-Paläontologisches Institut der 824 p.
Universität Hamburg, 86:81–188. LIPIARSKI, I. 1971. Dolnopermska flora martwicy karniowickiej koło
BARKER, G. M. 2001. Gastropods on land: phylogeny, diversity and Krakowa. Instytut Geologiczny, Prace, 58:5–80.
adaptative morphology, p. 1–146. In G. M. Barker (ed.), The Biology of LORIOL, P. de AND A. JACCARD. 1865. Etude géologique et paléontolo-
Terrestrial Molluscs. CAB International, Oxford. gique de la formation d’eau douce infracrétacée du Jura, et en
BATTEN, R. L. 1995. Pennsylvanian (Morrowan) Gastropods from the particulier de Villers-le-Lac. Mémoires de la Société de Physique et
Magdalena Formation of the Hueco Mountains, Texas. American d’Histoire naturelle de Genève, 18:1–68.
Museum Novitates, 3122:1–46. MARTINS, A. M. F. 1996. Anatomy and systematics of the Western Atlantic
BOLE, J. 1974. Rod Zospeum Bourguignat 1856 (Gastropoda, Ellobiidae) Ellobiidae (Gastropoda: Pulmonata). Malacologia, 37:163–332.
v Jugoslaviji. Razprave, Slovenska Akademija Znanosti in Umetnosti, NORDSIECK, H. 1986. The system of the Stylommatophora (Gastropoda),
17:251–291. with special regard to the systematic position of the Clausiliidae, II.
BOUCHET, P. AND J-P. ROCROI. 2005. Classification and Nomenclator of Importance of the shell and distribution. Archiv für Molluskenkunde,
Gastropod Families. Malacologia, 47:1–397. 117:93–116.
COX, L. R. 1926. Anthracopupa brittanica sp. nov., a Land Gastropod OPLUŠTIL, S. 2004. Late Carboniferous tectono-sedimentary evolution
from the Red Beds of the Uppermost Coal-Measures of Northern and related terrestrial biotic changes on the North Variscan and
Worcestershire. Quarterly Journal of the Geological Society, 82:401– Appalachian forelands, and adjacent paralic and continental basins.
410. Geologica Balcanica, 34:51–67.
PACYNA, G. 2003. Carboniferous fructifications from a new locality in
CUVIER, G. L. C. 1797. Tableau Èlèmentaire de l’Histoire Naturelle des
Sosnowiec – Preliminary report. Proceedings XXVI Symposium
Animaux [des Mollusques]. Baudonin, Paris, 710 p.
Geology of Coal-bearing strata of Poland, Cracow, 121–125.
ĆWIŻEWICZ, M. AND J. SZULC. 1989. Warunki klimatyczne środowiska
PACYNA, G. AND D. ZDEBSKA. 2001. Upper Carboniferous plant
sedymentacji martwicy karniowickiej. Przegla˛d Geologiczny, 37:180–
macrofossils from sideritic concretions of the Upper Silesian Coal
187.
Basin of Sosnowiec. Preliminary report. Materiały XXIV Sympozjum
DAWSON, J. W. 1855. Acadian geology; an account of the geological Geologia formacji we˛glonośnych Polski, Kraków, 75–79.
structure and mineral resources of Nova Scotia and portions of the PACYNA, G. AND D. ZDEBSKA. 2002. Upper Carboniferous plant
neighboring provinces of British America. Oliver and Boyd, Edinburgh, macrofossils from sideritic concretions in Sosnowiec (Upper Silesian
388 p. Coal Basin) and Mazon Creek (Illinois, USA). Proceedings, XXV
DAWSON, J. W. 1867. On the discovery of a new pulmonate mollusk Symposium Geology of Coal-bearing strata of Poland, Cracow, 123–127.
(Zonites (Conulus) priscus, Cpr.) in the coal-formation of Nova Scotia, PAN, H. 1982. Late Triassic-Early Jurassic Gastropods from Eastern
with a description of the species by Philip P. Carpenter. Quarterly Hunan and Northeastern Guangxi. Memoirs of Nanjing Institute of
Journal of the Geological Society of London, 23:330–333. Geology and Paleontology, 17:85–112.
DAYRAT, B. AND S. TILLIER. 2002. Evolutionary relationships of PANOW, E. 1936. Permokarbońska fauna martwicy karniowickiej.
euthyneuran gastropods (Mollusca): a cladistic re-evaluation of Rocznik Polskiego Towarzystwa Geologicznego, 12:36–41.
morphological characters. Zoological Journal of the Linnean Society, PFEIFFER, L. 1854. Synopsis Auriculaceorum. Malakozoolologische
135:403–470. Blätter, 1:145–156.
DAYRAT, B., A. TILLIER, G. LECOINTRE, AND S. TILLIER. 2001. New PILSBRY, H. A. 1900. On the zoological position of Partula and
Clades of Euthyneuran Gastropods (Mollusca) from 28S rRNA Achatinella. Proceedings of the Academy of Natural Sciences of
Sequences. Molecular Phylogenetics and Evolution, 19:225–235. Philadelphia, 52:561–567.
DESHAYES, G. P. (1861–1864). Description des Animaux sans vertèbres du RACIBORSKI, M. 1891. Permokarbońska flora karniowickiego wapienia.
Bassin de Paris, 2. Paris. Rozprawy Akademii Umieje˛tności, Wydział matematyczno-przyrod-
FALCON-LANG, H. J., M. J. BENTON, S. J. BRADDY, AND S. J. DAVIES. niczy, Ser. 2, 1:353–394.
2006. The Pennsylvanian tropical biome reconstructed from the Joggins SANDBERGER, F. (1870–1875). Die Land- und Süsswasser-conchylien der
Formation of Nova Scotia, Canada. Journal of the Geological Society, Vorwelt. C. W. Kreidel’s Verlag, Wiesbaden. 1000 p.
London, 163:561–576. SOLEM, A. AND E. L. YOCHELSON. 1979. North American Paleozoic Land
FILIPIAK, P. AND P. KRAWCZYŃSKI. 1996. Westphalian xiphosurans Snails, With a Summary of Other Paleozoic Nonmarine Snails. U.S.
(Chelicerata) from the Upper Silesia Coal Basin of Sosnowiec, Poland. Geological Survey Professional Paper, 1072:1–42.
Acta Palaeontologica Polonica, 41:413–425. STWORZEWICZ, E. 1999a. Miocene land snails from Bełchatów (Central
FISCHER, P. 1885. Description d’une nouvelle espèce de Dendropupa, du Poland), IV: Pupilloidea (Gastropoda Pulmonata). Systematic, bio-
terrain Permien de Saône-et-Loire. Journal de Conchyliologie, 33:99– stratigraphic and paleoecological studies. Folia Malacologica, 7:133–
105. 170.
GERBER, J. 1996. Revision der Gattung Vallonia Risso 1826 (Mollusca: STWORZEWICZ, E. 1999b. Miocene land snails from Bełchatów (Central
Gastropoda: Valloniidae). Schriften zur Malakozoologie aus dem Haus Poland), III. Carychiinae (Gastropoda; Pulmonata: Ellobiidae). Pa-
der Natur – Cismar, 8:1–227. läontologische Zeitschrift, 73:261–276.
GOLDBERG, S., J. MA, AND J. O’DONOHUE. 2003. Mass Extinction. The SZULC, J. AND M. ĆWIŻEWICZ. 1989. The Lower Permian freshwater
Traprock, 2:5–18. carbonates of the Slawków Graben, Southern Poland: sedimentary
HASZPRUNAR, G. AND G. HUBER. 1990. On the central nervous system of facies context and stable isotope study. Paleogeography, Paleoclima-
Smeagolidae and Rhodopidae, two families questionably allied with the tology, Paleoecology, 70:107–120.
Gymnomorpha (Gastropoda: Euthyneura). Journal of Zoology, TILLIER, S., M. MASSELOT, AND A. TILLIER. 1995 [1996]. Phylogenetic
220:185–199. relationships of the pulmonate gastropods from rRNA sequences, and
HEBERT, B. L. AND J. H. CALDER. 2004. On the discovery of a unique tempo and age of the stylommatophoran radiation, p. 267–284. In J. D.
terrestrial faunal assemblage in the classic Pennsylvanian section at Taylor (ed.) Origin and evolutionary radiation of the Mollusca. Oxford
Joggins, Nova Scotia. Canadian Journal of Earth Science, 41:247–254. University Press.
KLUSSMANN-KOLB, A., A. DINAPOLI, K. KUHN, B. STREIT, AND C. WADE, C. M., P. B. MORDAN, AND B. CLARKE. 2000. A phylogeny of the
ALBRECHT. 2008. From sea to land and beyond – New insights into the land snails (Gastropoda: Pulmonata). Proceedings of the Royal Society
evolution of euthyneuran Gastropoda (Mollusca). BMC Evolutionary of London, 268:413–422.
Biology, 8:57. WADE, C. M., P. B. MORDAN, AND F. NAGGS. 2006. Evolutionary
KNIGHT, J. B., R. L. BATTEN, E. L. YOCHELSON, AND L. R. COX. 1960. relationships among the Pulmonate land snails and slugs (Pulmonata,
Paleozoic and some Mesozoic Caenogastropoda and Opistobranchia, p. Stylommatophora). Biological Journal of the Linnean Society, 87:593–
1310–1324. In R.C. Moore (ed.) Treatise on Invertebrate Paleontology, 610.
Part I, Mollusca 1. Geological Society of America, Inc. and University WENZ, W. 1923. Gastropoda extramarina tertiaria, p. 1069–1420. In C.
of Kansas Press, Lawrence. 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– YEN, T. C. 1949. Review of Paleozoic non-marine Gastropods and a
480. In O. H. Schindewolf (ed.) Handbuch der Paläozoologie, Band 6. description of a new genus from the Carboniferous rocks of Scotland.
Borntraeger, Berlin. Journal of Molluscan Studies, 27:235–240.
WHITFIELD, R. P. 1881. Notice of a new genus and species of air- ZARE˛CZNY, S. 1894. Mapa geologiczna okolic Krakowa i Chrzanowa.
breathing mollusk from the Coal-Measures of Ohio, and observations Kraków, 280 p.
on Dawsonella. American Journal of Sciences, ser. 3, 21, 125–
128. ACCEPTED 12 JULY 2009