Annales Societatis Geologorum Poloniae (2013), vol. 83: 179–200.

SYS TEM ATIC AND PALAEO ECO LOGI CAL STUDY OF MIOCENE TERRES TRIAL GASTRO PODS FROM ZWIERZYNIEC (SOUTHERN POLAND)

Ewa STWORZEWICZ1, Valentin A. PRISYAZHNYUK2 & Marcin GÓRKA3

1 Insti tute of System at ics and Evolu tion of Ani mals, Polish Academy of Sciences, S³awkowska 17, PL 31-016 Kraków, Poland; e-mail: [email protected] 2 In sti tute of Geo log i cal Sci ences, Ukrai nian Acad emy of Sci ences, 55b Gonchara, UK 01030 Kiev, Ukraine; e-mail: [email protected] 3 Faculty of Geol ogy, Univer sity of Warsaw, Al. ¯wirki i Wigury 93, PL 02-089 Warszawa, Poland; e-mail: [email protected]

Stworzewicz, E., Prisyazhnyuk, V. A. & Górka, M. 2013. System atic and palaeo eco logi cal study of Mio cene ter res trial gas tro pods from Zwierzyniec (south ern Po land). Annales Societatis Geologorum Poloniae, 83: 179–200.

Ab stract: While the ma rine molluscs from Sarmatian de posits of the Carpathian Foredeep Ba sin are known in general, there is a paucity of data on the terres trial gas tropods. Recently, a rich assem blage of terres trial snails, ac compa nied by fresh wa ter spe cies, was found in Zwierzyniec, in the north-west ern, mar ginal part of the Carpa- thian Foredeep. Among the 38 taxa recog nised, there are 22 spe cies found in Po land for the first time; a new clau- siliid spe cies Triloba magurkai Stworzewicz sp. nov. is de scribed. Fresh wa ter gas tro pods (nine Lymnaeidae spe - cies and two Planorbidae species) were presented elsewhere. The malacofauna com prises aquatic and typ ical hygrophilous el e ments from coastal wet land hab i tats, some xerophilous spe cies from dry, open en vi ron ments, and gas tro pods from an ad ja cent sub trop i cal wood land.

Key words: ter res trial gas tro pods, Mid dle Mio cene, Carpathian Foredeep Ba sin, sys tem at ics, palaeo ec ol ogy.

Manu script re ceived 25 October 2013, ac cepted 16 December 2013

IN TRO DUC TION

Dur ing the Early Sarmatian, the Carpathian Foredeep thors have suggested an im portant role for storm-gener ated Basin was the northern most part of the Paratethys. The ba- rip-cur rents in the trans port of de tri tal mate rial (Rutkowski, sin was filled mainly by a thick se quence of clayey de pos its 1976; Czapowski, 1984; Czapowski and Studencka, 1990). (Krakovets (Krakowiec) Clay of Machów Forma tion; see A recent study (Górka, 2003) points to torren tial rains as the Œliwiñski et al., 2012), but its northern part was domi nated main factor of transport, causing rapid washout and flooding by lit to ral fa cies, commonly called the “de tri tal Sarmatian” of the adjoin ing land areas. As a re sult, some gas tropod as- and repre sent ing the formal lithostratigraphic unit, named semblages were found in sev eral lo cali ties of this area, but the Chmielnik Forma tion (see Alexandrowicz et al., 1982). that from Zwierzyniec is the most di versi fied. Further subdi vi sion is based on lithostratigraphy and this The strati graphic posi tion of the sequence un der discus - distinguishes three in for mal lithostratigraphic units within sion has remained un clear for a long time. While the forami- the “de trital Sarmatian” se quence. These are called “ho ri - nifer as semblages helped to de ter mine the age as low er most zons” (Rutkowski, 1976) or “com plexes” (Górka, 2003, Sarmatian Anomalinoides dividens Zone (£uczkowska and 2008b). The lower complex consists of calcar e ous sandsto- Rutkowski, 1970), results based on nannofossils were less nes and conglomerates and occurs only locally. The middle defin i tive, rang ing from NN5 through NN5/NN6 to NN6 part is com posed mainly of fine sands with thin inter cala tions zones (Dudziak and £aptaœ, 1991). The most useful data of clays and micritic limestones. The upper complex is the seem to be those obtained from the analy sis of in situ mol - most di verse, includ ing clays, sands/sand stones and coarse lus can as semblages. The re sults in di cate that both the “de - gravels/con glom er ates of var i ous or i gins. The main source of trital Sarmatian” and the un der lying clays of the Machów clastic ma terial, trans ported into the Sarmatian sea, was the Formation in the area dis cussed rep re sent the Lower Sar- ad join ing land ar eas, com posed of var ied Me so zoic rocks matian (i.e. Volhynian; see Czapowski and Studencka, and older Mio cene (mainly Badenian) depos its. Some au- 1990; Studencka, in press). 180 E. STWORZEWICZ ET AL.

Fig. 1. Palaeo geo graphi cal and geo log i cal back ground. A. Poland within palaeogeographic frame work of the Paratethys realm. B, C. Geo log i cal sketch-maps of the Carpathian Foredeep and vi cin ity of Busko-Zdrój, re spec tively. D. Lo ca tion of the gas tro pod-bear ing site at Zwierzyniec (mod i fied af ter Górka, 2003)

GEO GRAPH ICAL AND GEO LOG I CAL are abundant shells of Plicatiforma praeplicata (Hilber), SET TING Mactra (Sarmatimactra) eichwaldi Laskarew, Ervilia dis- sita Eichwald, E. trigonula Sokolov, Loripes niveus (Eich- Zwierzyniec is sit u ated in the north-west ern, mar ginal wald). The thickness of the unit exposed is 1.2 m. part of the Carpathian Foredeep (Fig. 1); its dis tance from Unit II. Fine-grained, grey quartz sand with hori zon tal the land dur ing the Sarmatian time can be es timated at some lami na tion at the base of the unit, becom ing deformed 10 kilometres (Zastawniak, 1980). The local ity is an aban- and/or homog e nous upward. The upper part of the unit doned sand-pit, situ ated in a densely forested gorge, about 4 comprises car bon ate con cre tions, form less in ser tions and km north of Busko-Zdrój, close to the Kraków – Chmielnik lenses of pelitic lime stone of lac us trine or i gin. Ma rine road (20°46´E, 50°32´N). The sandy succes sion at Zwierzy- molluscs are pres ent as in layer 1, but ter res trial and (rarely) niec was first de scribed by Friedberg (1928). He gave a de- fresh wa ter snails (Anisus, Lymnaea) prevail. Other terres - tailed de scrip tion of the pro file, with spe cial at ten tion to the trial ele ments are small frag ments of wood and rare rep tile pres ence of the marine Sarmatian fauna. bones. The thickness of the unit varies from 0.2 to 0.7 m. A rich terres trial gastro pod fauna, found in the Zwie- Unit III. Fine-grained, hor i zon tally lam i nated quartz rzyniec sec tion, co mes al most ex clu sively from de pos its of sand with thin inter ca la tions of sandy marls of lagoonal or the middle com plex, repre sented by fine-grained sands with lac ustrine ori gin. The upper part of the unit, com posed of thin in ter ca la tions of micritic limestones. The sec tion is as car bon ate con cre tions, con tains some ma rine gas tro pods follows (Fig. 2). and bivalves. The thickness varies between 0.6 and 0.7 m. Unit I. Light grey quartz sand, fine- to coarse-grained, Unit IV. Fine-grained, light brown, grey ish, lo cally with thin in ter ca la tions of pelitic lime stone. Hor i zon tal lam- yellowish quartz and carbon ate sands. This unit is hori zon - i na tion is pres ent, but nu mer ous synsedimentary de for ma- tally lami nated at the base, but is homo ge neous or with de- tions are domi nant. There are two hori zons of coarse sand formed laminae at the top (0.25 m). There are marine mo- with ripple-cross strat ifi ca tion in the topmost part of the lluscs sim ilar to those in layer 1, but there are also nu merous unit. The faunal assem blage is repre sented by frequent small gastro pods of the Hydrobiidae and Rissooidea. Fresh- Foraminifera and remnants; among the lat ter there wa ter, stagnophilous gas tro pods (Lymnaea, Anisus) and MIO CENE TER RES TRIAL GAS TRO PODS FROM ZWIERZYNIEC 181

Fig. 2. Litho-strati graphi cal back ground. A. Syn thetic profile of Mio cene de posits of the study area (lithostratigraphic units af ter Alexandrowicz et al., 1982). B. Profile of the “detri tal Sarmatian”. C. Lithological pro file of the Zwierzyniec site with depositional struc- tures. Strati graphic po sition of the shell-bear ing layer is in di cated by an ar row crushed shells of ter res trial snails (Cepaea sp.) occur rarely. or “complex I” (sensu Czapowski and Studencka, 1990). At The bed thickness is 0.65 m. the Zwierzyniec site, it is observed in the upper most part of Unit V. Fine-grained sands with marly in ter ca la tions the pro file, in the west ern part of the gorge. As may be pre- sim ilar to those in layer 3. The fauna is extremely poor and sumed from ob ser va tions at ad ja cent lo cal i ties (see Cza- com posed of frag ments of marine mollusc shells. The unit powski and Studencka, 1990), this unit lies di rectly on top thickness is 0.4 m. of “middle complex” with an erosional boundary. Unit VI. Fine-grained, homog e nous quartz sands with numer ous limy concrections. The shape of concrections is usually irreg u lar and their dimen sions vary from a few to 30 MA TE RIAL AND METH ODS cm. The thickness is esti mated as 2–2.5 m. Unit VII. Fine-grained con glom er ates, built pre dom i- Most of the mate rial was collected by M. Górka during nantly of lime stone clasts with a small ad mixture of quartz his fieldwork, over a long period in the 1980s and at the be - and chert grains. Within strongly lithified hori zons, an in- ginning of the 1990s, comple mented by the first two authors distinct lam ina tion can be ob served. The base of the unit is in 1996. The shells and their identi fi able fragments were ob- not exposed, and the thickness exceeds 1.5 m. tained through the sieving of sandy sedi ments on a sieve of Units I to VI repre sent the “middle hori zon” or “middle 0.25 mm mesh and some of the large-sized shells were taken com plex” (sensu Rutkowski, 1976; Górka, 2003, respecti- di rectly from the sed iment. The mate rial was not rich, and vely) or “complex I” (sensu Czapowski and Studencka, 1990). only shells of the small-sized spe cies were nu mer ous and The pres ence of bi valve as semblage – Plicatiforma praepli- well-preserved, while zonitid and helicid shells were dam- cata (Hilber), Mactra (Sarmatimactra) eichwaldi Laskarew, aged and less numer ous, with the excep tion of Cepaea Ervilia dissita Eichwald, E. trigonula Sokolov, Loripes niveus shells. Fresh wa ter snails (9 spe cies of Lymnaeidae and 2 (Eichwald) – deter mines the age of this se quence as the low er - spe cies of Planorbidae) were desribed ear lier (Prisyazhnyuk most part of the Early Sarmatian (i.e. early Volhynian). et al., 2006). Unit VII repre sents the “upper hori zon” or “up per com- All of the measure ments were taken with a Nikon mea- plex” (sensu Rutkowski, 1976; Górka, 2003, respec tively) surescope MM-11, accu racy 0.01 mm. The fol lowing ab- 182 E. STWORZEWICZ ET AL.

Localities breviations in the text below denote shell param e ters: H – Taxa shell height, B – shell breadth, h – aper ture height, b – aper - Zwierzyniec Be³chatów Opole ture breadth, HLW – last whorl height, NW – number of Microstele ikvae Prisyazhnyuk * whorls. Microstele buryaki Steklov * The clas sifi ca tion of the above the fam ily Granaria antiqua (Schübler) * level fol lows Bouchet and Rocroi (2005). Most of the mate - Granaria sp. * rial is kept at the In sti tute of Sys temat ics and Evo lu tion of Ani mals, Polish Academy of Sciences in Kraków, under acuminata (Klein) * * Gastrocopta nouletiana (Dupuy) collection numbers Ml/1035– Ml/1070. Only Cepaea and * * sensu lato Pseudochloritis shells are in the col lec tion of the Fac ulty of Ge ol ogy, War saw Uni ver sity, ab bre vi ated ZC C1–ZC C23 Gastrocopta serotina Ložek * and ZC H1–ZC H4. Gastrocopta fissidens (Sandberger) * *? Gastrocopta ex. gr. didymodus * (Sandberger) RE SULTS Gastrocopta 2 species 3 species gottschicki Wenz * Thirty eight taxa of terres trial snails were iden tified in joossi (Gottschick) ** the sed iments from Zwierzyniec (Ta ble 1). Twenty two of Strobilops 5 species 2 species them were new records for Po land. The re maining are subcyclophorella ** known either from Be³chatów (Stworzewicz, 1995, 1999a, Gottschick 1999b; Stworzewicz and So³tys, 1996; Stworzewicz and callosa (Reuss) * * * Prisyazhnyuk, 1997) or from Opole (Andreae, 1902, 1904), Vertigo protracta (Sandberger) * * and some from both lo cal ities. The clausilid Triloba ma- Vertigo angulifera O. Boettger * * gurkai Stworzewicz is de scribed here as new. Such a great Vertigo 3 species 2 species propor tion of taxa, pre viously un known in Poland, in the Zwierzyniec sed i ments is not sur pris ing, con sid er ing the Negulus suturalis (Sandberger) * * * cir cum stances of their de po si tion, dif fer ent from those at Serrulastra trolli Nordsieck * Triloba magurkai Stworzewicz Be³chatów and Opole, and as so ciated with the Paratethys * shore. An un iden ti fied spe cies of Myosotella, re garded as n. sp. one of the “supratidal and estuarine ellobiids” (Morton, Clausiliidae 9 species 1955), is an indi ca tor of mari time condi tions. Pupilloids are Discus costatus (Gottschick) * the most fre quent, par tic u larly Gastrocopta, which is repre - Helicodiscus roemeri (Andreae) * * * sented by five species and more than a hundred speci mens. Hawaiia antiqua Riedel * Phenacolimax suevica (Sandberger) * Ta ble 1 Vitrea procrystallina Andreae * * Miozonites costatus (Sandberger) * Gastro pod taxa found in Zwierzyniec and their occur rence Aegopinella cf. subnitens (Klein) * in the other Polish Mio cene lo cal ities Klikia giengensis (Klein) * (the remain ing species from Be³chatów and Opole are not Psudochloritis incrassatus (Klein) * included, only the number of species from genera found Cepaea sylvestrina gottschicki * also in Zwierzyniec are re corded) Wenz

Localities Taxa Zwierzyniec Be³chatów Opole Pomatias rivulare (Eichwald) ** SYS TEM ATIC PALAE ON TOL OGY Carychium starobogatovi Steklov * Carychium schlckumi schlickumi Clade LITTORINIMORPHA Golikov and Starobogatov, * Strauch 1975 Carychium sp. * Family POMATIIDAE Newton, 1891 Ge nus Pomatias Studer, 1789 Carychium 10 species 4 species Type spe cies: Nerita elegans Müller, 1774 Myosotella sp. * aff. paramonovae * Prisyazhnyuk Pomatias rivulare (Eichwald, 1829) Azeca sp. * Fig. 3A Azeca 2 species *1829 Cyclostoma rivularis sp. nov. – Eichwald: p. 302. steinheimensis (Miller) * 1926 Pomatias rivulare (Eichwald) – Lindholm, p. 167. 1966 Pomatias rivulare (Eichwald) – Steklov, p. 121, pl. 1, Pupilla sp. ex gr. iratiana * figs 8–10. Microstele caucasica Steklov * Ma te rial: Two spec i mens and several shell frag ments (Ml/1035). MIO CENE TER RES TRIAL GAS TRO PODS FROM ZWIERZYNIEC 183

Di men sions: H = 9.6 mm, 12.35 mm; B = 7.32 mm, 10.06 mm; h De scrip tion: The shell frag ment is very thin and fine with a shiny = 4.36 mm, 6.16 mm; b = 4.33 mm, 5.95 mm; HLW = 7.15 mm, sur face and in dis tinct ra dial lines. Un der higher mag ni fi ca tion, 9.89 mm; NW = 4.5, 4.75. fine pricks are visi ble, form ing spi ral lines. Be low the su ture, a Re marks: The smaller shell is strikingly simi lar to those of P. paral lel groove is marked and some verti cal, thin, comma-like cav- rivulare from Be³chatów (Be³-B, Middle Mio cene) and from sev- i ties be low the groove, which are most prob a bly hair traces. The eral Mio-Pliocene local ities in Ciscaucasia (Steklov, 1966). The ap er ture, nar rowly drop-shaped, is sharp ened in its upper part and larger one is somewhat more elon gated and the sur face sculp ture is rounded at the bot tom. Its margin is simple in the up per half and more distinct. Such spec i mens were also found at Be³chatów thick ened by a cal lus on the in ner side of the bot tom half. There (Stworzewicz, 1995). are two lamellate teeth in the aper ture. The columellar is situ ated on the columellar-basal transi tion and con tin ues as an ar cu ate Clade J. Morton, 1955 lamella, deflected upward around the . The pari etal tooth Family ELLOBIIDAE L. Pfeiffer, 1854 is stout, situ ated close to the columellar margin. Both teeth en ter Subfamily CARYCHIINAE Jeffreys, 1830 deeply in side the body whorl (about 0.75 of whorl). Re marks: The speci men from Zwierzyniec was tenta tively as- Ge nus Carychium O. F. Müller, 1774 signed to Myosotella be cause among the gen era of Ellobiidae only Type spe cies: Carychium min i mum Müller, 1774 some Myosotella mem bers share most shell features with the shell frag ment discussed. These are the out line of the last whorl and ap - Carychium starobogatovi Steklov, 1966 er ture, teeth, and above all, the microsculpture, con sisting of spi ral Fig. 3B rows of pits and one spi ral row of hair traces. Ellobiids of some other gen era may have hir sute whorls, but then the shape of the *1966 Carychium starobogatovi sp. nov. – Steklov: p. 124, fig. shell and ap er ture and the dentition are dif fer ent (Mar tins, 1996). 40, pl. 2, fig. 14. No hair traces were described in fos sil rep resen ta tives of the Ma te rial: Two spec i mens and one damaged shell (Ml/1036). Ellobiidae (Wenz, 1920; Zilch, 1959–1960; Kókay, 2006). Di men sions: H = 1.67 mm, 1.8 mm; B = 0.69 mm, 0.73 mm; h = 0.57 In its shell form and dentition, the shell fragment is most simi lar mm, 0.63 mm; b = 0.52 mm, 0.53 mm; HLW = 0.97 mm, 1.00 mm. to Stoliodoma (Stoliodomopsis) glandina (Boettger, 1874) from Re marks: The two shells are sim i lar in outline and the weak de - the Mid dle Oligocene, but the latter species is twice as large and it vel op ment of ap er ture bar ri ers to the spec i mens, de scribed by has no spi ral struc ture or spi ral groove be low the su ture (Wenz, Steklov (1966) from the Mio-Plio cene of Ciscaucasia. How ever 1920). On the other hand, such sur face sculp ture of spi rally ar - one of them is dis tinctly larger than all of the other known shells. ranged small pits was de scribed by Kókay (2006, p. 50) in Auri- culastra badeniensis Kókay, 2006, from the Up per Badenian coal- Carychium schlickumi schlickumi Strauch, 1977 bear ing clay at Pusztamiske (Hun gary), but no spi ral groove be low Fig. 3C the su ture or whorl traces were men tioned. Un for tu nately, the di - mensions of the spec i mens from Pusztamiske were not presented, *1977 Carychium schlickumi schlickumi ssp. nov. – Strauch: p. and the holotype is somewhat dif fer ent in shape (Kókay, 2006, pl. 168, pl. 16, figs 40–47, pl. 19, figs 68–70, 72–73, 75. 16, figs 10–11). Ma te rial: Eight spec i mens (ML/1037). This is the first re cord of land ellobiids in Po land, other than that Di men sions: H = 1.75–2.05 mm; B = 0.88–0.96 mm; h = 0.70– of Carychiinae. 0.77 mm; b = 0.56–0.64 mm; HLW = 1.06–1.2 mm. Re marks: The spec i mens from Zwierzyniec do not dif fer in the Clade Schmidt, 1855 general shell outline and struc ture of the inter nal lamellae from Family Pilsbry, 1900 those, known from the Plio cene of Ger many (Strauch, 1977) and from the Upper Mio cene of Ukraine (Steklov, 1966, p. 127, fig. Ge nus Azeca Fleming, 1828 42, as Carychium suevicum O. Boettger), but they are gen er ally Type spe cies: He lix goodalli Férussac, 1821 some what larger. Azeca aff. paramonovae Prisyazhnyuk, 1990 (in Luleva Carychium sp. and Prisyazhnyuk, 1990) Ma te rial: One spec i men (Ml/1038). Fig. 3E Di men sions: H = 1.9 mm; B = 0.86 mm; h = 0.72 mm; b = 0.63 *1990 Azeca paramonovae sp. nov. – Prisyazhnyuk (in Luleva mm; HLW = 1.12 mm. and Prisyazhnyuk): p. 36, text-fig. Re marks: The shell outline resem bles C. nouleti, but the structure Ma te rial: One spec i men and three frag ments of the last whorl of the inter nal lamellae is simi lar to that of C. antiquum. The lack with an ap er ture (Ml/1040). of suf fi cient ma te rial pre cludes the pos si bil ity of any spe cific as- Di men sions: H = 4.44 mm; B = 1.83–2.09 mm; h = 1.35–1.59 sign ment. mm; b = 1.24–1.38 mm; HLW = 2.45–2.79 mm. De scrip tion: The only well-pre served shell con sists of 6 whorls. Subfamily PYTHIINAE Odhner, 1925 The ap er ture mar gin of all the spec i mens is surrounded by a Ge nus Myosotella Monterosato, 1906 cord-like edge. There are four teeth in the ap er ture, vis i ble in front Type spe cies: Auricula myo so tis Draparnaud, 1801 view. A long, lamellate pari etal tooth is divided into two parts by an in ci sion. Its outer part is shorter than the in ner one, reach ing Myosotella sp. more then 0.5 whorl, deep in side the shell. Of the two columellar Fig. 3D lamellate teeth, the lower is situ ated close to the aper ture margin, whereas the up per one, deeply set and poorly visi ble in front view, Ma te rial: One fragment of shell: the last whorl with aper ture as cends ver ti cally on the columella to ca. 0.3 of the body whorl (Ml/1039). height. The pala tal tooth is sit uated in the mid dle of the pala tal Di men sions: B = 1.94 mm; h = 2.04 mm; b = 1.23 mm; HLW = wall and does not reach the ap er ture mar gin; it is roll-like or di - 2.87 mm. vided by an in ci sion into two knobs. Be sides, two elon gate teeth 184 E. STWORZEWICZ ET AL. MIO CENE TER RES TRIAL GAS TRO PODS FROM ZWIERZYNIEC 185

are placed very deeply in the body whorl, at a distance of ca. 0.5 Some shells also have a small infrapalatal tooth. whorl from the ap er ture mar gin and they are in visi ble in front Re marks: The speci mens from Zwierzyniec were identi fied as re- view. fer able to P. steinheimensis from the up per Mio cene de posits at Re marks: The speci mens from Zwierzyniec are very distinct, be- Steinheim, despite some slight dif ferences in ap ertural char ac ters. cause of the lower columellar tooth. In this respect, they are most The two spec i mens described by Miller (1900) have a somewhat sim i lar to Azeca paramonovae, described from the Lower Sarma- stouter ap er ture mar gin and columellar tooth, but the scanty mate - tian of Volhynia (Luleva and Prisyazhnyuk, 1990, p. 36). More - rial from both lo cal i ties does not per mit the ex clu sion of intraspe - over, they are both fusiform and the only dif fer ence is in the pal a - cific varia tion. From among the few sinistral Pupilla species tal tooth, which is very weakly marked in the spec i mens from known from the Neogene, only P. belokrysi Steklov, 1966 seems Zwierzyniec and well-de vel oped in those from Volhynia. to be very simi lar in its shell form and size, as well as in the contin - The only Azeca spe cies, pre vi ously de scribed from Poland, is ua tion of the columellar tooth, shown by Prisyazhnyuk (2008a), Azeca frechi Andreae, 1902 from Opole (Andreae, 1902, p. 14), which is partic u larly char ac teris tic of older (Karaganian) forms of which has a hor i zon tal columellar tooth, an un di vided pa ri etal one, this spe cies. How ever, P. belokrysi has a somewhat less-devel - and a distinctly smaller shell. oped ap er ture mar gin. All other sinistral spesies are much smaller – P. blainvilleana Azeca sp. (Dupuy, 1850) or big ger – P. rathi (Sandberger, 1858), P. muta- Fig. 3F bilis Steklov, 1966 and P. staszicii £omnicki, 1886; they dif fer in the num ber of whorls, and the struc ture of the columellar tooth is Ma te rial: Two frag ments of last whorl with somewhat dam aged un known. ap er ture (pa ri etal tooth bro ken) (Ml/1041). Di men sions: B = 1.84 mm, 1.90 mm; h = 1.47 mm, 1.53 mm; b = 1.24 mm, 1.38 mm; HLW = 2.51 mm, 2.67 mm. Pupilla ex gr. iratiana (Dupuy, 1850) Re marks: The spec i mens differ from the pre ced ing spe cies in the Fig. 3H form of the columellar teeth, par tic u larly the lower one, which is knob-shaped. The tooth on the pal a tal wall is ab sent, but there is a Ma te rial: Two spec i mens (one of them damaged) and eight frag - tooth on the pal a tal mar gin of ap er ture. In ad di tion, the cord-like ments with ap er ture (Ml/1043). edge along the ap er ture mar gin is weakly marked. Di men sions: H = 2.86 mm, 2.52 mm; B = 1.61 mm, 1.54 mm; h = 0.96 mm, 0.84 mm; b = 0.83 mm, 0.83 mm; HLW = 1.44 mm, 1.46 mm. Family Turton, 1831 De scrip tion: Oval-cy lindri cal shells consist of six moder ately Ge nus Pupilla Leach in Turton, 1831 con vex whorls, cov ered by weakly marked, blunt, un evenly spa- Type spe cies: Pupa marginata Draparnaud, 1801 ced radial striae. The body whorl ascends be fore the ap erture. The crest is clearly marked and thick ened to ward the um bi li cus, partly Pupillla steinheimensis (Miller, 1900) cov er ing it. It is fol lowed by a deep groove be hind the lip. The ap - Fig. 3G er ture mar gins are slightly re flexed and joined by a thin or very thin pa ri etal cal lus. *1900 Pupa (Pupilla) steinheimensis sp. nov. – Miller: p. 398, There are 4–5 teeth in the aper ture. The lamellate pari etal tooth pl. 7, fig. 15. reaches ca. 0.4 whorl deep in side the shell. The an gu lar tooth is Ma te rial: Two spec i mens and five frag ments includ ing the body weakly de vel oped in the form of a small knob, rarely elon gated, whorl with ap er ture (Ml/1042). but most fre quently this is not per cep ti ble. The columellar tooth is Di men sions: H = 2.66 mm, 2.60 mm; B = 1.47 mm, 1.56 mm; h = well-devel oped, broadened at the base. The two pala tal teeth do 0.82 mm, 0.77 mm; b = 0.80 mm, 0.78 mm; HLW = 1.00 mm, 1.21 not reach the aper ture margin and the lamellate lower pala tal tooth mm. is lon ger than the up per, which is usu ally knob-shaped. De scrip tion: Sinistral, oval-cy lindri cal shells consist of 6.5–7 Re marks: Pupilla sp. from Zwierzyniec cor re sponds very closely whorls, weakly con vex and sep a rated by moder ately deep su tures. to the de scrip tion and il lus tra tion of P. iratiana by Dupuy (1850, The body whorl is somewhat nar rower than the two above and it p. 310, pl. 15, fig. 7) and to the more de tailed redescription and ascends before the aper ture. The surface sculpture is composed of draw ings by Bourguignat (1881, p. 65, pl. 3, figs 83–85). How - ir reg u lar, del i cate striae, which are al most in vis i ble. The crest is ever, it dif fers from the P. iratiana spec i mens, pre sented by Finger strongly marked, fol lowed by a deep groove behind the lip and (1998, pl. 12, fig. A) and by Harzhauser et al. (2008, p. 49, figs paral lel to it. At the base, the crest passes into a basal keel. The ap- 5.1–3), in the less distinct shell sculp ture, stouter shell shape and erture is semicir cu lar, with weakly reflexed mar gins and thin lip, more cir cu lar ap er ture. There is no in for mation in Fin ger’s pa per joined by a thin pari etal cal lus. The follow ing teeth are always about the dentition of P. iratiana. On the other hand, the spec i men pres ent: lamellate pa ri etal tooth; weakly de vel oped an gu lar tooth, pre sented by Harzhauser et al. (2008) has only one small pal a tal which is most often in the form of a low nodule; deeply sit uated tooth. Some other sim i lar forms were de scribed, i.e. Pupilla ira- columellar tooth, in vis i ble in ap er tural view or marked as a small tiana eumeces Boettger, 1884, Pupilla iratiana max ima Boettger, nod ule, con tin ues in side the body whorl as a ridge and spi rally as - 1908 and Pupilla iratiana suevica Gottschick and Wenz, 1919, but cends the columella. There is also a short lamellate pala tal tooth. their tax o nom i cal po si tion still re mains am big u ous.

Fig. 3. Land gas tro pods from Zwierzyniec; scale bars 1 mm. A. Pomatias rivulare (Eichwald, 1829), Ml/1035. B. Carychium starobogatovi Steklov, 1966, Ml/1036; C. Carychium schlickumi schlickumi Strauch, 1977, Ml/1037; D. Myosotella sp., Ml/1039; E. Azeca aff. paramonovae Prisyazhnyuk, 1990 (in Luleva and Prisyazhnyuk, 1990), Ml/1040; F. Azeca sp., Ml/1041; G. Pupillla steinheimensis (Miller, 1900), Ml/1042; H. Pupilla ex gr. iratiana (Dupuy, 1850), Ml/1043; I. Microstele caucasica Steklov, 1966, Ml/1044; J. Microstele ikvae Prisyazhnyuk, 1971, Ml/1045; K. Microstele buryaki Steklov, 1966, Ml/1046; L. Granaria antiqua (Schübler in Zieten, 1830), Ml/1047; M. G. antiqua, teeth on pala tal wall, Ml/1047a; N. Granaria sp., Ml/1048 186 E. STWORZEWICZ ET AL.

Ge nus Microstele Boettger, 1886 Ta ble 2 Type spe cies: Pupa moltei O. Boettger, 1886 Among the few Microstele speci mens from Zwierzyniec, repre - Di men sions of Microstele buryaki Steklov, 1966 (in mm) senting three species, there is a marked trend towards weaken ing of the pari etal, columellar and pala tal teeth. Only the an gular tooth H 3.73 3.87 3.89 3.96 4.09 re mains con stant. Apart from M. caucasica, M. buryaki and M. B 1.81 1.88 1.83 1.85 1.82 ikvae, there are two shell frag ments with only an an gu lar tooth h 1.32 1.41 1.40 1.43 1.50 present, while the remain ing teeth are entirely re duced. They may b 1.21 1.34 1.28 1.25 1.30 rep re sent an ini tial stage of tran sition to the younger spe cies M. alamellata Steklov, 1966 which re sembles but the ma te - HLW 2.05 2.15 2.01 2.08 2.13 rial from Zwierzyniec is insuf fi cient to identify them. Microstele has not been found pre vi ously in Poland. De scrip tion: Elon gated, ta per ing shells con sist of 6 whorls, cov - ered with very fine, irreg u lar striae. The aper ture has a reflected margin and a some what thick ened lip. In the ap er ture, only the an - Microstele caucasica Steklov, 1966 gular tooth is well-de vel oped, whereas the pa ri etal and columellar Fig. 3I teeth are very small or vesti gial and deeply sit uated. There is no pal a tal tooth. *1966 Microstele caucasica sp. nov. – Steklov: p. 160, pl. 4, Re marks: The dentition of spec i mens from Zwierzyniec is very figs 84–85. sim ilar to that of speci mens from the Mid dle Mio cene (Karagan) Ma te rial: Three fragments contain ing last whorl with well-pre- of Ciscaucasia (Steklov, 1966); how ever, the lat ter are some what served ap er ture (Ml/1044). big ger and less elon gated. Di men sions: B = 1.85–1.89 mm; h = 1.34–1.38 mm; b = 1.29– 1.27 mm; HLW = 2.06–2.14 mm. Family Steenberg, 1925 De scrip tion: The sur face of the last whorl is very finely and ir reg - u larly stri ated. The umbi li cus is very nar row, but open. The ap er - Ge nus Granaria Held, 1837 ture has a reflexed and thickened margin. There are 4 teeth in the Type spe cies: Pupa frumentum Draparnaud, 1801 ap er ture: an gu lar, deeply sit u ated pa ri etal and pal a tal, and colu- mellar. Granaria antiqua (Schübler in Zieten, 1830) Re marks: Re gard ing the num ber and form of the teeth, the spec i - Fig. 3L, M mens from Zwierzyniec are very simi lar to both M. wenzi (Fischer, 1920) and M. caucasica. However, the shell breadth and ap er ture *1830 Pupa antiqua sp. nov. – Schübler in Zieten: p. 39, pl. 29, di mensions show that it is M. caucasica, which is clearly smaller fig. 7a–c. than the pre ced ing spe cies. 1846 Pupa Schübleri n. nov. – Klein, p. 74, pl. 1, fig. 18. 1916 Torquilla antiqua (Zieten) – Gottschick and Wenz, p. 61. Microstele ikvae Prisyazhnyuk, 1971 1923c antiqua antiqua (Zieten) – Wenz, p. 940. Fig. 3J Ma te rial: Three spec i mens (one somewhat de formed) and 23 frag ments con tain ing the last whorl with an ap er ture (Ml/1047). *1971 Microstele ikvae sp. nov. – Prisyazhnyuk: p. 601, fig. Di men sions:(of two complete shells and four well-pre served frag - 1w, fig. 2.3. ments): H = 7.95 mm, 8.0 mm; B = 2.98–3.29; h = 2.29–2.55 mm; Ma te rial: Two spec i mens and two frag ments of the last whorl b = 1.96–2.35 mm; H/B = 2.65, 2.66; HLW = 3.2–3.63 mm. with an ap er ture (Ml/1045). De scrip tion: Shells consist of 8.5 almost flat whorls; the embry - Di men sions: H = 3.95 mm, 3.64 mm; B = 1.91 mm, 1.80 mm; h = onic whorls are smooth, the next seem to be smooth, but they are 1.46 mm, 1.48 mm; b = 1.38 mm, 1.20 mm; HLW = 2.29 mm, 2.23 cov ered with poorly visi ble reg ular striae, which are more distinct mm. on the last whorl before the ap er ture. There are 8–9 teeth in the ap - De scrip tion: Ta per ing shells con sist of 5.5–6 whorls, very weakly er ture. The arched angularis is thick ened or dou bled in front by a and ir reg u larly stri ated. The umbi li cus is open and deep. The ap er - small, knob-shaped tooth (subangularis) and joins the ap er ture ture mar gin is re flexed, with a weakly thick ened lip. In the ap er - margin; it reaches inside as far as the ante rior part of the deeply sit - ture, only an gular and pala tal teeth are well-devel oped; the pari etal uated pari etal tooth. In some shells there is a short lamellate tooth is very weak and deeply placed. The columellar tooth is ves ti gial, (spiralis) sit uated be yond the in ner end of the angu lar lamella. The in the form of a hardly marked knob, in visi ble in front view, or ab - pari etal tooth is high lamellate and thick, situ ated in the mid dle sent. part of the pari etal wall, and reaches ca. half-whorl deep inside the Re marks: The close compar i son of spec i mens from Zwierzyniec shell. The columellar and subcolumellar teeth are deeply set; the to those from the Lower Sarmatian de posits of Volhynia-Podolia for mer is thicker and longer than the lat ter. There are four teeth on does not raise any doubt that they are conspecific, al though the the pala tal wall; two long lamellate teeth (lower pala tal and upper former shells are some what smaller. pal a tal) reach the in ner edge of the lip and be come higher within but only the lower pala tal tooth is strongly de vel oped and al ways clearly visi ble (Fig. 3M). The remain ing two (infrapalatal and Microstele buryaki Steklov, 1966 suprapalatal) are deeply sit uated and short-lamellate or knob- Fig. 3K shaped. In some spec i mens, the infrapalatal tooth has a hardly marked con tin u a tion to the in ner lip mar gin. *1966 Microstele buryaki sp. nov. – Steklov: p. 161, fig. 58, pl. Re marks: The spec i mens from Zwierzyniec cor re spond to the fig - 4, figs 86–87. ure and short de scrip tion of Pupa antiqua from the Upper Mio cene Ma te rial: Seven spec i mens (includ ing two partly damaged) and de posits at Steinheim am Aalbuch (Schübler in Zieten, 1830) and three frag ments with an ap er ture (Ml/1046). to the de scrip tion of Pupa schuebleri (Klein, 1846), re garded by Di men sions: Given in in Ta ble 2. some authors as the valid name of the former species (Miller, MIO CENE TER RES TRIAL GAS TRO PODS FROM ZWIERZYNIEC 187

1900; Gottschick and Wenz, 1916) or as a sep a rate spe cies (Harz- One spec i men dif fers from the oth ers in hav ing a swol len shell hauser and Binder, 2004). How ever, Klein (1846) wrongly in ter - and less con vex whorls. Such a form was de scribed from Sansan as preted the name Pupa antiqua as pre-oc cu pied, ar gu ing that Ma- G. lartetii (Dupuy, 1850, p. 307) and subse quently was re garded theron (1832) gave the same name to an other spe cies (sim i lar to as the subspe cies lartetii of G. acuminata (Wenz, 1923c, p. 919). He lix tridens Müller, 1774 = Pupa tridens, see Draparnaud, 1801), In its shell shape and the form of parieto-angu lar and columellar be cause he took no ac count of the fact that Matheron’s work was teeth, it is also simi lar to some speci mens of G. acuminata lartetii published two years later (Schübler in Zieten, 1830; Matheron, from Ukraine (Brykov, Zamiehov, Bogdanovskij Karier), al - 1832). though in Gozhik and Prisyazhnyuk (1978, p. 89, pl. 11, figs 10, On the other hand, the spec i mens from Zwierzyniec dif fer from 14), spec i mens with basal tooth are pre sented. those, mis iden ti fied by Lueger (1981) as G. antiqua (see Harz- G. acuminata is one of the most wide spread and the longest hauser and Binder, 2004). occuring spe cies in west ern and cen tral Eu rope, known from the In its shell form and dentition, G. antiqua from Zwierzyniec is Middle Mio cene to the Middle-Late Pliocene (Manganelli and sim i lar to G. subvariabilis (Sandberger), known from the Up per Giusti, 2000). Oligocene and the Lower Mio cene (Sandberger, 1858), but it is some what smaller and more distinctly stri ated. The H/B ratio for spec i mens from Hochheim is 2.52 (Boettger, 1889); for two spec i - Gastrocopta nouletiana (Dupuy, 1850) sensu lato mens from Zwierzyniec it is 2.65 and 2.66, and the lower pal a tal Fig. 4B tooth is the most de vel oped. On the other hand, the teeth, par tic u - *1850 Pupa Nouletiana sp. nov. – Dupuy: p. 309, pl. 15, fig. 6. larly on the pala tal wall, are the same in G. antiqua and Recent G. Ma te rial: 35 spec i mens and some fragments with an ap er ture variabilis (Draparnaud, 1801). Ac cord ing to Boettger (1889), (Ml/1050). those three spe cies make an evo lu tion ary con tin uum subvaria- Re marks: Three mor pho log i cal forms of this spe cies, wide spread bilis–antiqua–variabilis. in Eu rope, were found at Zwierzyniec. They all com monly oc cur This is the first Miocene record for Granaria in Poland. in the Mid dle-Up per Mio cene de posits and are also known from Poland (Stworzewicz, 1999b). Granaria sp. 1. The typ ical form, as a rule ovate with three pala tal teeth, corre - Fig. 3N sponds to that de scribed by Dupuy (1950) from Sansan, that found in Be³chatów (Stworzewicz, 1999b, p. 163–164, figs 59, 61) and Ma te rial: One frag ment of last whorl with ap er ture and ap er tural that from Steinheim (Gottschick and Wenz, 1919, p. 12, pl. 1, figs bar ri ers (Ml/1048). 22–23). Di men sions: B = >2.70 mm; h = 2.59 mm; b = 2.28 mm; HLW = Di men sions: See Table 3. 3.5 mm. Re marks: The spec i men dif fers from those of G. antiqua in: 1. the Ta ble 3 distinctly, regu larly stri ated surface of the last whorl; 2. the dis- tinctly thick ened ap er ture mar gin; 3. the subangular tooth dis- Di men sions of Gastrocopta nouletiana (Dupuy, 1850) tinctly sepa rated from the an gular; 4. the infrapalatal and upper sensu lato, the typ ical forms (in mm) pala tal teeth, al most as long and stout as the lower pala tal; and 5. the pres ence of a third columellar lamella above the upper colu- H 2.27 2.05 2.12 1.91 2.15 mellar on the dor sal side (in visi ble in front view). B 1.37 1.27 1.33 1.31 1.33 The speci men is most simi lar to the Recent G. frumentum (Dra- parnaud, 1801) and may be re garded as its an ces tral form. The ma- h 0.86 0.81 0.75 0.68 0.80 te rial is insuf fi cient for a descrip tion of it as a new species. b 0.87 0.74 0.80 0.73 0.75 HLW 1.41 1.25 1.28 1.22 1.24 Family Pilsbry, 1918 H/B 1.66 1.61 1.60 1.46 1.62 Ge nus Gastrocopta Wollaston, 1878 Type spe cies: Pupa acarus Benson, 1856 2.Ovate-elon gated form with two pal a tal teeth; the lower one is solid and ar cu ate. Gastrocopta acuminata (Klein, 1846) Di men sions: See Table 4. Fig. 4A Ta ble 4 *1846 Pupa acuminata sp. nov. – Klein: p. 75, pl. 1, fig. 19. Di men sions of Gastrocopta nouletiana (Dupuy, 1850) Ma te rial: Three spec i mens (includ ing the larger and swol len one, sensu lato, ovate-elongated forms (in mm) re garded as G. acuminata larteti) and four frag ments of the last whorl with an ap er ture (Ml/1049). H 2.27 2.12 2.27 2.14 2.40 Di men sions: H = 2.60–2.95 mm, 3.01 mm; B = 1.60–1.75 mm, 1.92 mm; h = 0.94–1.07 mm, 1.08 mm; b = 0.84–0.98 mm, 1.08 B 1.31 1.27 1.30 1.30 1.27 mm; HLW = 1.45–1.64 mm, 1.84 mm. h 0.86 0.85 0.76 0.75 0.86 Re marks: The speci mens from Zwierzyniec differ somewhat b 0.84 0.77 0.72 0.77 0.78 from the typi cal form of G. acuminata, pre sented in Gottschick HLW 1.37 1.28 1.24 1.25 1.27 and Wenz (1916, p. 62–64, pl. 1, fig. 4) and Schlickum (1976, p. H/B 1.73 1.67 1.75 1.65 1.89 10, pl. 1, fig. 26), in hav ing the interpalatal knob-shaped tooth. On the other hand, the interpalatal tooth is of ten ob served in the spec i - mens from Ciscaucasia (Steklov, 1966). The speci mens from 3. An elon gated form, usu ally re garded as G. nouletiana graci- Zwierzyniec are within the variabil ity range of G. acuminata from lidens, with three or four pala tal teeth, but the interpalatal tooth is Be³chatów as well; some of the over 300 speci mens found there al ways present (Stworzewicz, 1999b, fig. 60). have a small interpalatal tooth (Stworzewicz, 1999b). Di men sions: See Table 5. 188 E. STWORZEWICZ ET AL.

Ta ble 5 the outer surface of the pala tal wall, there are two dark streaks, cor re sponding in po sition to the two pal a tal teeth, but there is no Di men sions of Gastrocopta nouletiana (Dupuy, 1850) crest. The umbi li cus is open, but nar row and deep. There are 7 sensu lato, elongated forms (in mm) teeth in the ap er ture. The parieto-an gu lar tooth con sists of two sep- a rate, paral lel parts, arcuately bent outward. The pari etal portion H 2.50 2.40 2.28 2.16 2.32 con sider ably over laps the an gular por tion (half a length, or even more). A small infraparietal tooth is pres ent in all the speci mens B 1.42 1.38 1.40 1.28 1.30 from Zwierzyniec. The columellar tooth is subhorizontal and the h 0.82 0.73 0.80 0.85 0.83 basal is short lamellate or knob-shaped. There are three teeth on b 0.82 0.76 0.72 0.80 0.76 the pala tal wall; the upper one (suprapalatal tooth) is small or HLW 1.41 1.25 1.28 1.22 1.24 barely marked. Re marks: The speci mens from Zwierzyniec corre spond gener ally H/B 1.76 1.74 1.63 1.68 1.78 to those de scribed and il lustrated by Sandberger (1863, p. 57, pl. 5, figs 16a–c), Wenz (in Fischer and Wenz, 1914, p. 97, pl. 6, figs Gastrocopta serotina Ložek, 1964 19a–b) and Wenz (1921, pl.18, figs 20–21). In deed, there is no Fig. 4C infraparietal tooth in the G. fissidens fig ures of Wenz, but it is pres ent in Sandberger’s fig ures and in Boettger’s (1889, p. 286) re- *1964 Gastrocopta serotina sp. nov. – Ložek: p. 194, figs 1–4. marks on the G. fissidens spec i mens from Mosbach-Biebrig and Ma te rial: 36 well pre served spec i mens and some fragments with Wiesbaden (Lower Miocene). It is also distinctly marked in G. an ap er ture (Ml/1051). fissidens infrapontica, de scribed by Wenz from Loebersdorf (Wenz, Di men sions: H = 1.66–1.73 mm and 1.90–2.42 mm; B = 1.00– 1927, p. 47, pl. 2, fig. 8) and af ter wards pre sented by Lueger (1981, 1.07 mm and 1.10–1.29 mm; h = 0.54–0.64 mm and 0.66–0.84 p. 27, pl. 2, figs 20–21). The shells from Zwierzyniec come closest mm; b = 0.52–0.59 mm and 0.61–0.81 mm; HLW = 0.97–1.05 mm in shape and ap er tural bar ri ers to the lat ter form, but also to three and 1.04–1.37 mm. spec i mens of G. fissidens from Hochheim, stored at the Forschung- Re marks: The speci mens from Zwierzyniec are identi cal in their sinstitut und Naturmuseum Senckenberg (no. 152 289/3). Ac cord - shell shape and aper tural bar ri ers to those de scribed by Ložek ing to Wenz, G. fissidens infrapontica is smaller (H = 1.7 mm; B = (1964) from the de posits of Ctinìves (Cen tral Bo he mia). How- 0.9 mm) than typi cal G. fissidens and has more con vex whorls, but ever, they vary in size (hav ing the same num ber of whorls, with an the spec i men pre sented by Lueger (1981) is slightly big ger (H = av er age of 5) enough to distin guish two groups (see Di mensions). 1.8 mm) and falls within the vari abil ity range of spec i mens from The larger shells pre domi nate and they also have some what more Zwierzyniec (see de tailed mea sure ments). Wenz’s di ag no sis mas sive ap er tural bar ri ers. “Diese Form unterscheidet sich von Gastrocopta (Sinalbinula) fissidens (Sandb) lediglich durch etwas geringere Größe und we- niger hohe, etwas stärker gewölbte Umgänge“ is based on one spe- Gastrocopta fissidens (Sandberger, 1863) cimen only. Thus, the form described may rep re sent one ex treme Fig. 4D of the vari abil ity range of the nom i nal form. *1863 Pupa fissidens sp. nov. – Sandberger: p. 57, pl. 5, figs G. fissidens has been re corded from Poland be fore as Pupa 16a–c. (Vertigo) didymodus var. fissidens Sandberger, 1874 found in the 1875 Pupa (Ver tigo) didymodus var. fissidens Sandberger – Sarmatian sand de posits from Sobów near Tarnobrzeg (Friedberg, Sandberger, p. 399. 1905), a lo cal ity sit u ated not far from Zwierzyniec. 1889 Leucochilus fissidens (Sandberger) – Boettger, p. 285. More over, the shells from the de posits of Be³chatów, iden ti fied 1914 Leucochila fissidens (Sandberger) – Fischer and Wenz, as G. cf. ferdinandi (Andreae, 1902) in Stworzewicz (1999b), are p. 97, pl. 6, fig. 19. ac tu ally very large (H of 30 spec i mens is 1.83–2.40 mm) and some what more slen der spec i mens of G. fissidens. Ma te rial: 22 spec i mens and some fragments with ap er ture (Ml/ 1052). Di men sions (of 22 spec i mens): H = 1.72–2.07 mm; B = 0.96– Gastrocopta ex gr. didymodus (Sandberger, 1858) 1.11 mm; h = 0.62–0.74 mm; b = 0.61–0.70 mm; HLW = 1.02– Fig. 4E 1.18 mm; H/B = 1.79–1.90. De tailed mea sure ments (in mm) of 10 spec i mens show a contin - Ma te rial: Ten spec i mens and five frag ments with ap er ture (Ml/ u ous tran sition be tween small and large forms (Ta ble 6). 1053). Di men sions: H = 1.95–2.14 mm; B = 0.92–1.07 mm; h = 0.67– Ta ble 6 0.77 mm; b = 0.63–0.72 mm; HLW = 1.08–1.24 mm; H/B = 2.0–2.18. Di men sions of Gastrocopta fissidens (Sandberger, 1863) De scrip tion: Oval-cy lin dri cal shells con sist of 5–5.25 moder ately (in mm) con vex whorls, sep a rated by a moder ately deep su ture. The sur- face is finely, but distinctly stri ated. On the outer surface of the H 1.72 1.78 1.85 1.89 1.93 1.95 1.97 2.00 2.04 2.07 body whorl, there is a distinct crest, paral lel to the aper ture margin. There are 6–7 teeth in the ap er ture. The parieto-an gu lar tooth con - B 0.96 1.03 1.00 1.03 1.06 1.05 1.06 1.07 1.08 1.11 sists of two parts, but they are not paral lel (in contra dis tinc tion to h 0.62 0.63 0.64 0.71 0.70 0.70 0.70 0.72 0.74 0.70 those in G. fissidens), be cause the pari etal part only slightly over - b 0.61 0.61 0.63 0.67 0.65 0.66 0.62 0.68 0.67 0.64 laps the an gular part. Most fre quently, the an te rior part of the pa ri - HLW 1.04 1.02 1.08 1.07 1.09 1.10 1.09 1.15 1.16 1.13 etal tooth only reaches the poste rior part of the small an gu lar tooth. The infraparietal tooth is present in some speci mens. The remain - H/B 1.79 1.83 1.85 1.83 1.82 1.86 1.86 1.87 1.88 1.86 ing teeth, namely the columellar, basal and palat als, are as in G. fissidens. De scrip tion: Ovate-turreted shells con sist of 4.75–5 con vex Re marks: At first, Sandberger (1858–1863) de scribed both G. whorls, very finely stri ated and sep a rated by a dis tinct su ture. On didymodus and G. fissidens but later (1870–75) he was un cer tain, MIO CENE TER RES TRIAL GAS TRO PODS FROM ZWIERZYNIEC 189

as to whether they were two sep a rate species. Af terwards, Boet- Ma te rial: 15 spec i mens and 35 frag ments of the last whorl with an tger (1889) on the ba sis of the orig i nal ma te rial from Hochheim ap er ture and the in ner lamellae vis i ble (Ml/1055). showed some impor tant interspecific differ ences. First of all, the Di men sions: H = 1.32–1.77 mm; B = 1.93–2.33 mm; h = 0.58– parieto-an gu lar tooth in the for mer species is not “zweispaltige” 0.79 mm; b = 0.99–1.22 mm; HLW = 0.84–1.14 mm; B/H = like in G. fissidens but “zweizipfelige” (see also descrip tions of the 1.32–1.54; NW = 4.75–5.25. two species from Zwierzyniec and Figs 4D, E). G. didymodus dif - De scrip tion: The shell is widely coni form, with somewhat con vex fers from G. fissidens also in its clearly more cy lindri cal shell, less whorls, sep a rated by a moder ately deep su ture. The embry onic convex whorls, and distinctly marked crest. These charac ters are whorls (1.75) are smooth and the oth ers are sculp tured, with very clearly visi ble in the speci mens from Zwierzyniec, but G. didy- distinct ribs (40–55 on the last whorl), which are somewhat wea- mo dus is known only from the Upper Oligocene/Lower Mio cene ker but clearly visi ble at the base. The last whorl is more or less an- de posits, which are signif i cantly older than those in Zwierzyniec. gled at the pe riph ery. The um bi li cus is open, but very nar row, and It is prob a ble that the spec i mens from Zwierzyniec be long to a it expands abruptly, just be fore the ap erture. The pari etal lamella is new species, being a direct descen dant of G. didymodus. How ever, higher and thicker than the infraparietal, and in con trast to the lat - this view requires compar a tive studies of all species, regarded as ter, it reaches the end of pari etal callus. The infraparietal lamella members of the didymodus group, i.e. G. obstructa (Sandberger, rises in side the ap er ture and grad u ally co mes level with the pa ri - 1874), G. obstructa francofurtana (Boettger, 1877), G. baudoni etal. They both ex tend half way round the last whorl. The edges of (Michaud, 1862) and G. farcimen (Sandberger, 1874). these lamellae are very weakly crenate, if at all. The interparietal lamella is absent. The columellar lamella is very weakly marked Family Steenberg, 1925 or absent. There are three basal plicae, arranged in a sequence Ge nus Leiostyla Lowe, 1852 from the short est one near the columella, to the longest; in some spec i mens the third plica is equal to the sec ond. The mid dle one is Type spe cies: Leiostyla vincta Lowe, 1852 the most massive and the highest; gener ally it is rather vari able. Re marks: The speci mens from Zwierzyniec are very close to Leiostyla cf. gottschicki (Wenz, 1922) those from Be³chatów, iden ti fied as Strobilops cf. joossii (Gott- schick) in Stworzewicz (1999b, figs 41–43). More ma te rial has es - *1922 gottschicki sp. nov. – Wenz: p. 107, text-fig. tablished that the distinc tion among the three somewhat sim i lar Ma te rial: One frag ment of the last whorl with partly damaged ap - spe cies, i.e. S. joossii, S. costata (Clessin, 1877) and S. tiarula erture and a splin ter of pari etal callus with pari etal and angu lar (Sandberger, 1886) is clear. The latter species is inter me di ate in teeth (Ml/1054). charac ter between the two former. Di men sions: (of ap er ture): h = 1.06 mm; b = 1.08 mm. The dif fer ences be tween the remain ing strobilopsids were noted De scrip tion: The surface of the last whorl is finely striated. The by Manganelli et al. (2008), who re viewed the known Eu ro pean aper ture margin is moder ately re flexed, with a somewhat thick- Neo gene spe cies, on the ba sis of re search on strobilopsids from ened lip. The um bi li cus is open, but nar row and slit-like. Only one Balze di Capreno. columellar tooth and two pala tal teeth are visi ble in the preserved fragment. The two teeth, visi ble on the splinter of the pari etal wall, con tinue in side the last whorl. The deeply en ter ing columellar Family Morse, 1864 tooth be gins on the lip and runs obliquely downwards, then bends Ge nus Vallonia Risso, 1826 upwards, pro longed as a large, thin lamella, paral lel to the colu- Type spe cies: He lix pulchella Müller, 1774 mella. The lower pala tal tooth is long, rather low near the aper ture, and be coming higher within. There is a small sec ond pal a tal tooth Vallonia subcyclophorella Gottschick, 1911 above, but as the ap er ture mar gin is partly broken, its form is hard Fig. 4H, I to identify. Re marks: From among a few species known from the Neo gene, *1911 He lix (Vallonia) subcyclophorella sp. nov. – Gott- only L. gottschicki de scribed from Steinheim (Wenz, 1922) and schick: p. 503, pl. 7, figs 2–2b. also noted from Hollabrunn (Schütt, 1967) has such a com pli cated Ma te rial: Ten spec i mens and five dam aged shells (Ml/1056). columellar tooth with the verti cal part deep within the aper ture. Di men sions (in brack ets, those of a clearly bigger spec imen): H = The frag ment from Zwierzyniec con tains only the body whorl with 0.88–1.05 (1.24) mm; B = 1.86–2.35 (2.67) mm; h = 0.76–0.9 an ap er ture, so the shape and size of the shell re main un known. (1.09) mm; b = 0.76–0.98 (1.17) mm; B umbi li cus = 0.52–0.71 Because of the com plicated columellar tooth, L. gottschicki comes (0.86) mm; B protoconch = 0.50–0.63 (0.66) mm; NW = 3.0–3.6 closest to the Recent Cauca sian species L. caucasica (Pfeif fer, (3.75). 1857) and L. paulinae (Lindholm, 1913, pre sented by Schileyko, Re marks: The form of the shell and aper ture, as well as the sur- 1984). face striation of speci mens from Zwierzyniec, fit very well with V. subcyclophorella from numer ous other sites. The umbi li cus of Family Wenz, 1915 some speci mens is ec cen tric, but ac cord ing to Gerber (1996), this Strobilops Pilsbry, 1892 is not un usual in the spe cies. V. subcyclophorella was widely dis - trib uted in the Mio cene of Eu rope, and it is also known from Type spe cies: He lix labyrinthica Say, 1817 Kazakhstan (Steklov and Tsytovich, 1967).

Strobilops joossi (Gottschick, 1911) Family Fitzinger, 1833 Fig. 4F, G Ge nus Ver tigo Müller, 1774 *1911 Strobilus Joossii sp. nov. – Gottschick: p. 502, pl. 7. figs Type spe cies: Ver tigo pusilla Müller, 1774 16a–c. 1915 Strobilops joossi (Gottschick) – Wenz, p. 80, fig. 9, pl. 4, Ver tigo callosa (Reuss, 1849) figs 14a–c. Fig. 4J 1999b Strobilops cf. joossi (Gottschick) – Stworzewicz, p. 154, figs 41–43. *1849 Pupa callosa sp. nov. – Reuss: p. 30, pl. 3, fig. 7. 190 E. STWORZEWICZ ET AL. MIO CENE TER RES TRIAL GAS TRO PODS FROM ZWIERZYNIEC 191

Ma te rial: One spec i men (Ml/1057). Di men sions: H = 1.7, 1.74 mm; B = 0.81, 0.86 mm; h = 0.63, Di men sions: H = 2.33 mm; B = 1.45 m; h = 0.94 mm; b = 0.80 0.65 mm; b = 0.51, 0.54 mm; HLW = 1.08, 1.15 mm. mm; HLW = 1.38 mm; NW = 5.25. Re marks: The two spec i mens do not dif fer from those, known De scrip tion: The shell is ovate with poorly con vex whorls. There from sev eral Neo gene lo cal i ties in Eu rope, in clud ing Be³chatów are 6 teeth in the aper ture: angu lar, pari etal, columellar, basal and and Opole (Stworzewicz, 1999b). 2 pala tal. The most char ac teris tic feature is the long and thin lamellate an gular tooth, located closer to the pa rietal callus margin Family CLAUSILIIDAE Gray, 1855 than the pa ri etal tooth, which is par al lel to the for mer one. The Ge nus Serrulastra Nordsieck, 1981 columellar tooth is short, lamellate, deeply and obliquely situ ated, with the in ner part turned up ward. Type spe cies: Clausilia amphiodon Reuss, 1860 Re marks: Because of the shell shape and size, the speci men from Zwierzyniec cor re sponds to V. callosa var. max ima Boettger, 1877 Serrulastra trolli Nordsieck, 1981 from Steinheim (Gottschick and Wenz, 1919, p. 14, pl. 1, fig. 27; Fig. 5A Finger, 1998, pl. 9 fig. A), but dif fers from the lat ter in the form and lo ca tion of the an gu lar tooth and po si tion of the columellar *1981 Serrulastra trolli sp. nov. – Nordsieck: p. 76, pl. 8, fig. tooth. Al most identi cal spec i mens were found in the Lower Sar- 17. matian and the lower part of the Middle Sarmatian depos its of Ma te rial: Two frag ments of body whorl with partly pre served Volhynia-Podolia (Ukraine). lamellae and two fragments of pala tal wall with an aper ture (Ml/ 1061). Vertigo protracta (Sandberger, 1874) Di men sions: H fragm = 3.63 mm, 3.60 mm; h = 2.11 mm, 2.03 mm; b = 1.58 mm, 1.70 mm. *1874 Pupa protracta sp. nov. – Sandberger: p. 400. De scrip tion: The body whorl is cov ered with strong and reg u lar 1889 Ver tigo (Alaea) protracta (Sandberger) – Boettger, p. ribs, which are somewhat sparse on the dor sal side. The neck is 300, pl. 7, fig. 5. reg ularly rounded. The ap er ture is broadly oval, with a strongly Ma te rial: One spec i men and one frag ment (Ml/1058). folded peri stome. The su pe rior lamella is con nected with the spi ral Di men sions: H = 1.57 mm; B = 1.05 mm; h = 0.58 mm; b = 0.60 one, the in fe rior lamella steeply as cends and bends to the peri - mm; HLW = 0.97 mm. stome, forming a bi fur ca tion. In one spec i men, there is a small fold Re marks: The sin gle spec i men does not dif fer from those, known be tween the branches of the bi fur ca tion. The subcolumellar la- from other Neo gene lo cal i ties of Eu rope, in clud ing that at Be³- mella reaches the peristome. The clausilial appa ra tus con sists of chatów (Stworzewicz, 1999b). three pala tal plicae: a short up per plica (in one speci men it is con- nected with lunella), a more or less distinctly marked lunella, and a strong lower plica, which is vis i ble in side the ap er ture in front Ver tigo angulifera O. Boettger, 1884 view. The prin ci pal plica is very long and ends later ally. The clau- Fig. 4K silium is unknown. *1884 Ver tigo (Alaea) angulifera sp. nov. – O. Boettger: p. Re marks: The two clausiliid fragments are most simi lar to the 271, pl. 4, figs 10a–c. Serrulastra (Serruplica) spe cies, de scribed from the Mid dle Mio - cene sedi ments from St.Veit a.d. Triesting (Aus tria). Ma te rial: One spec i men and one frag ment (Ml/1059). The short up per plica is sepa rate from the lunella in only one Di men sions: H = 1.45 mm; B = 0.78 mm; h = 0.48 mm; b = 0.45 spec i men from Zwierzyniec and in both spec i mens the lunella is mm; HLW = 0.80 mm. not con nected with the lower plica. Re marks: Hav ing a cylin dri cal shape, the only spec i men from Zwierzyniec is most sim i lar to V. (Alaea) angulifera milleri Gott- schick et Wenz, 1919 from Steinheim (Sarmatian) and to some of Ge nus Triloba Vest, 1867 the V. angulifera spec i mens from Be³chatów (Stworzewicz, 1999b). Type spe cies: Clausilia sandrii Küster, 1847

Ge nus Negulus Boettger, 1889 Triloba magurkai Stworzewicz new spe cies Type spe cies: Pupa reinhardti Jickeli, 1874 Fig. 5B, C Ma te rial: One spec i men (holotype), con sisting of body whorl Negulus suturalis (Sandberger, 1858) with an ap er ture (ISEA PAS, no. Ml/1062). Fig. 4L Di men sions: H fragm = 8.68 mm; B = 5.45 mm; h = 5.15 mm; b = 4.40 mm. *1858 Pupa suturalis sp. nov. – Sandberger: p. 54, pl. 6, fig. 1. Type lo cal ity and type level: Zwierzyniec, near Busko (south ern 1889 Negulus lineolatus (Al. Br.) – O. Boettger, p. 269, pl. 6, Poland); fine-grained sand de posits of the Sarmatian (the up per fig. 8. part of the Mid dle Mio cene, MN 7+8, ca. 12.5 Ma). 1914 Negulus suturalis (Sandberger) – Fischer and Wenz, p. Et y mol ogy: Ded i cated to Marcin Górka, who dis cov ered the lo - 142, pl. 5, fig. 13. cality at Zwierzyniec (“magurka” is the acro nym of his name in Ma te rial: Two spec i mens (Ml/1060). his e-mail address).

Fig. 4. Land gas tro pods from Zwierzyniec; scale bars 1 mm. A. Gastrocopta acuminata (Klein, 1846), Ml/1049; B. Gastrocopta nouletiana (Dupuy, 1850), Ml/1050; C. Gastrocopta serotina Ložek, 1964, Ml/1051; D. Gastrocopta fissidens (Sandberger, 1863), Ml/1052; E. Gastrocopta ex gr. didymodus (Sandberger, 1858), Ml/1053; F, G. Strobilops joossi (Gottschick, 1911), Ml/1055, one shell with plicae inside last whorl vis i ble; H, I. Vallonia subcyclophorella Gottschick, 1911, Ml/1056; J. Ver tigo callosa (Reuss, 1849), Ml/1057; K. Ver tigo angulifera O. Boettger, 1884, Ml/1059; L. Negulus suturalis (Sandberger, 1858), Ml/1060 192 E. STWORZEWICZ ET AL. MIO CENE TER RES TRIAL GAS TRO PODS FROM ZWIERZYNIEC 193

De scrip tion: The body whorl is nar rowed to the ap er ture. Its sur - Ma te rial: Four spec i mens (includ ing 3 with bro ken last whorls) face is nearly smooth in front; on the dor sal side, there are weak, and frag ments of 19 spec i mens (Ml/1063). wrinkly striae, giving the im pres sion of a gran ulated structure. The Di men sions (in brack ets – of dam aged spec i mens): H = 2.43 mm neck is rounded. The ap er ture is obliquely pear-shaped, with a re - (1.55, 1.83); B = 4.93 mm (3.52, 3.86); h = 1.07 mm (0.96, 0.98); b flexed edge and a thick ened peri stome, also on the pa ri etal por tion, = 1.85 mm (1.30, 1.43); B um bi li cus = 1.41 mm (1.0, 1.1); NW = with out folds. The spi ral and up per lamellae are not con nected. 5.75 (4.75, 5). The spi ral lamella is rather weak, much weaker in the an terior part De scrip tion: The shell is discoidal with a low coni form spire, con - and the upper lamella is well-devel oped. They are situ ated at dif- sisting of six ra di ally ribbed whorls with blunt keel on the pe riph - fer ent dis tances from the columella, but they are not over lap ping. ery. The ribs become somewhat weaker at the base. There is not a The lower lamella as cends spirally and ends just before the peri- distinct microsculpture on the protoconch apart from hardly visi - stome with a bipar tite knot. The subcolumellar lamella is not visi - ble adsutural radial wrinkles, sim i lar to that described in Dis cus ble in front view. There is no lunella in the clausilial appa ra tus; pleuradrus (Bourguignat, 1881) by Harzhauser and Binder (2004). only pal a tal plicae are pres ent, of which four are vis i ble on the The um bi li cus is widely open, tak ing al most one-third of mi nor neck sur face (Fig. 5C). The prin ci pal plica is the lon gest, the up per shell di ame ter, so that all the whorls, in clud ing the nu cleus, are pala tal plica is clearly shorter, two mid dle pala tal plicae are the clearly visi ble. The aper ture is widely semilunar with a simple shortest (the up per one is very weakly de vel oped, so it is not visi - peri stome. ble from with out), and the lower pal a tal plica is lon ger than the Re marks: The speci mens from Zwierzyniec match those, de - two lat ter and ar cu ate. The clausilium is unknown. scribed from the Sarmatian depos its at Steinheim and stored in the Re marks: Re gard ing the di mensions of the sin gle frag ment of col lec tions of the Senckenberg Mu seum (151372/5) and the Mu - shell, it must belong to a rather large clausiliid species. The clausi- seum of Natu ral His tory in Lon don. Re cently, new ma te ri als, con - lial appa ra tus, with out lunella, con sisting of 4–5 pal a tal plicae, and tain ing D. costatus from the kleini lay ers of the Steinheim Ba sin, the upper lamella, sepa rated from spiral lamella, are charac teris tic were pre sented by Finger (1998). of Triloba ge nus (Allopiinae) (Vest, 1867). Un til quite re cently, The spe cies has not been re corded from Poland to date. Triloba species have not been noted as fossils, apart from the un - des cribed Triloba n. sp., re corded from the Up per Mio cene de pos- its at Nerezi, near Skopje in Mac e do nia (Nordsieck, 2007). In Family HELICODISCIDAE H. B. Baker, 1927 2012, Schnabel described that spe cies as Triloba pappi,on the ba - Ge nus Helicodiscus Morse, 1864 sis of mate rial, collected from Nerezi more than half of a century Type spe cies: He lix lineatus Say, 1817 ago by Adolf Papp. Triloba magurkai is simi lar to T. pappi with re gard to di men- Helicodiscus (Helicodiscus) roemeri (Andreae, 1902) sions of the ap er ture, the nar rowed form of the last whorl and the clausilial appa ra tus of plicae-type. However, it differs in the posi - Fig. 5G tion of the upper-spi ral lamellae (the upper lamella barely reaches *1830 He lix depressa sp. nov. – Eichwald: p. 215. the spi ral one, but they are not overlaping), in a well-de vel oped 1853 He lix depressa Eichwald – Eichwald, p. 300, pl. 11, figs princi pal lamella, in a differ ent ending of lower lamella (it is not 10a–d. bent up wards) and in the less reflexed edge of the ap er ture. 1902 Hyalinia (Gyralina) roemeri sp. nov. – Andreae, p. 9, T. pappi was found in the Up per Mio cene de posits and it was re - fig. 3. garded as “the first known fos sil spe cies of the Alopiinae from 1942 Gyralina roemeri (Andreae) – Wenz and Edlauer, p. 93, Southeast Eu rope” (Schnabel, 2012). The pres ent find dem on - pl. 4, fig. 12. strates that in the Mid dle Mio cene, the group reached much fur ther 1972 Helicodiscus eichwaldi sp. nov. – Prisyazhnyuk, p. 132, north wards. figs a–d. 1979 Helicodiscus (Helicodiscus) roemeri (Andreae) – Family DISCIDAE Thiele, 1931 Schlickum, p. 69, fig. 3. Genus Discus Fitzinger, 1883 1997 Helicodiscus (Helicodiscus) depressus (Eichwald) – Type spe cies: He lix ruderata Férussac, 1821 Stworzewicz and Prisyazhnyuk, p. 202, figs 1–4. Ma te rial: One subadult spec i men (Ml/1064). Dis cus costatus (Gottschick, 1911) Di men sions: H = 1.10 mm; B = 2.27 mm; B umbi li cus = 0.96 mm; Fig. 5D–F NW = 3. Re marks: The single speci men of this rare Neogene species is *1911 Patula (Charopa) costata sp. nov. – Gottschick: p. 501, iden ti cal to those from other sites in Po land (Opole, Be³chatów), pl. 7, figs 15a–c. Austria and Ukraine. Detailed data on H. roemeri are included in 1916 (Gonyodiscus) costata (Gottschick) – Gott- Stworzewicz and Prisyazhnyuk (1997), where the au thors pre - schick and Wenz, p. 29. sented it un der the name Helicodiscus depressus, the species de- 1920 Gonyodiscus costatus (Gottschick) – Gottschick, p. 40. scribed by Eichwald (1830) as He lix depressa from Ho³owczyñce 1998 Dis cus costatus (Gottschick) – Fin ger, p. 18, pl. 10, figs (Podolia in Ukraine). However, Dietrich Kadolsky (pers. info., D–F. 1997) drew to the au thors’ at ten tion that “…Eichwald’s name is an

Fig. 5. Land gas tro pods from Zwierzyniec; scale bars 1 mm. A. Serrulastra trolli Nordsieck, 1981, Ml/1061; B, C. Triloba magurkai Stworzewicz sp. nov., holotype, Ml/1062, fron tal and dor sal view; D–F. Dis cus costatus (Gottschick, 1911), Ml/1063; G. Helicodiscus (Helicodiscus) roemeri (Andreae, 1902), Ml/1064; H. Hawaiia antiqua Riedel, 1963, Ml/1065; I. Vitrea procrystallina Andreae, 1902, Ml/1067; J, K. Phenacolimax suevica (Sandberger, 1874), Ml/1066; L. Miozonites costatus (Sandberger, 1874), Ml/1068; M, N. Aegopinella cf. subnitens (Klein, 1853), Ml/1069 194 E. STWORZEWICZ ET AL.

ob jec tive ju nior hom onym of He lix depressa Montagu, 1803, Re marks: The subadult speci men from Zwierzyniec matches the Menke, 1828 and Grateloup, 1827. There fore, it has to be re placed de scrip tion of V. procrystallina from Opole, al though Andreae by Hyalinia roemeri Andreae 1902, the next old est available spe - (1902, fig. 4) showed a shell with a very nar row umbi li cus. On the cies group name and a subjec tive synonym…”, af ter wards placed other hand, the spec i men from Zwierzyniec does not dif fer from in Helicodiscus (Prisyazhnuk, 1972). those pre sented from Rákosd (Gaál, 1911) and Zwiefaltendorf (Schlickum, 1976). It is a very rare spe cies in Mio cene de posits. Family Cockerell, 1891 Ge nus Hawaiia Gude, 1911 Ge nus Miozonites Pfeffer, 1930 Type spe cies: He lix minuscula Binney, 1840 Type spe cies: He lix algiroides Reuss, 1849

Hawaiia antiqua Riedel, 1963 Miozonites costatus (Sandberger, 1874) Fig. 5H Fig. 5L *1963 Hawaiia antiqua sp. nov. – Riedel: p. 37, figs 9–14. *1874 Archaeozonites costatus sp. nov. – Sandberger: p. 604. 1966 Hawaiia antiqua Riedel – Steklov, p. 208, figs. 81–82, 1916 Zonites (Aegopis) costatus (Sandberger) – Gottschick pl. 8. figs 161–162. and Wenz, p. 21, pl. 1, figs 1a–c. 2008 Hawaiia antiqua Riedel – Prisyazhnyuk, 2008b p. 95, Ma te rial: One spec i men (somewhat dam aged) and nu merous pl. 3, fig. 22, pl. 5, figs 45–46. fragments (Ml/1068). Ma te rial: One subadult spec i men (Ml/1065). Di men sions: H = 19 mm; B = 24.5 mm; h > 9.5 mm; b = > 12 mm; Di men sions: H = 0.99 mm; B = 1.78 mm; h = 0.71 mm; b = 0.68 HLW = 14 mm; NW = 5. mm; NW = 3.5. Re marks: In shell form, size and surface sculpture, the speci men Re marks: The sin gle spec i men with a bro ken ap er ture does not from Zwierzyniec cor re sponds to spec i mens from the Silvana- dif fer from those de scribed by Riedel (1963) from the Cau ca sus. schichten and Hohenmemmingen (Gottschick and Wenz, 1916) Al though the shell sur face of Hawaiia spe cies has char ac ter is tic and from the Karpatian de posits of the Korneuburg Ba sin (Nie- radial lines, they are barely vis ible on some of Riedel’s type speci - derösterreich) (Binder, 2002).This is the first re cord of the spe cies mens, or in the speci men from Zwierzyniec. in Poland. H. antiqua is known from several Mid dle and Late Mio cene sites in the Cau ca sus (Steklov, 1966), Podolia (Gozhik and Prisya- Ge nus Aegopinella Lindholm, 1927 zhnyuk, 1978) and Ser bia (Prisyazhnyuk, 2008b). Type spe cies: He lix pura Alder, 1830 This is the first re cord for Po land. Aegopinella cf. subnitens (Klein, 1853) Family VITRINIDAE Fitzinger, 1833 Fig. 5M, N Ge nus Phenacolimax Stabile, 1859 Type spe cies: Helico-Limax major Férussac, 1807 *1853 He lix subnitens sp. nov. – Klein: p. 210, pl. 5, fig. 7. partim 1875 Hyalinia orbicularis sp. nov. – Sandberger, p. 603, pl. 29, fig. 28. Phenacolimax suevica (Sandberger, 1874) 1916 Hyalinia (Polita) subnitens (Klein) – Gottschick and Fig. 5J, K Wenz, p. 23. *1874 Vitrina suevica sp. nov. – Sandberger: p. 602, pl. 29, fig. 1923a Oxychilus subnitens subnitens (Klein) – Wenz, p. 282. 27. 1976 Aegopinella subnitens (Klein) – Schlickum, p. 12, pl. 3, 1916 Vitrina (Phenacolimax) suevica Sandberger – Gott- figs 39–40. schick and Wenz, p. 24. Ma te rial: 11 juv. and subadult spec i mens (2–3.75 whorls), in the Ma te rial: One spec i men (Ml/1066). most part damaged (Ml/1069). Di men sions: H = 1.16 mm; Bmax = 2.16 mm; Bmin = 1.67 mm; h Di men sions: See Table 7. = 0.92 mm; b = 1.33 mm; NW = 2. Ta ble 7 Re marks: The subadult spec i men is a lit tle dam aged in the ap er - tural part. Nev er the less, it does not dif fer from those, de scribed Di men sions of Aegopinella cf. subnitens (Klein, 1853) from Steinheim and known also from several sites in Ukraine (in mm) (Gozhik and Prisyazhnyuk, 1978). H 2.57 2.43 2.05 1.97 - Family Mörch, 1864 B 5.03 4.80 4.09 3.75 3.79 Ge nus Vitrea Fitzinger, 1833 h 1.89 1.68 1.65 1.50 - Type spe cies: Glischrus (He lix) diaphanus Studer, 1820 b 2.29 2.22 1.77 1.69 - Bprot 1.12 1.05 1.08 1.04 1.11 Vitrea procrystallina Andreae, 1902 Bumb 0.94 - 0.89 0.82 - Fig. 5I NW 3.75 3.75 3.25 ca.3 ca.3 *1902 Hyalina (Vitrea) procrystallina sp. nov. – Andreae: p. 10, fig. 4. De scrip tion: The two big gest shells have 3.75 somewhat raised 1920 Vitrea (Vitrea) procrystallina Andreae – Gottschick, p. whorls, cov ered with weak, ir reg ular spaced growth lines, and 37. much less visi ble spi ral lines, with the ex cep tion of the smooth Ma te rial: One spec i men (Ml/1067). protoconch. On the basal side, this sculp ture is very weakly mar- Di men sions: H = 0.81 mm; B = 1.59 mm; h = 0.65 mm; b = 0.76 ked. The umbi li cus is open, all whorls are visi ble within; it takes mm; NW = 3.5. up ca. 1/5 of the shell di ame ter. The ap er ture is ovate-semilunar. MIO CENE TER RES TRIAL GAS TRO PODS FROM ZWIERZYNIEC 195

Re marks: The pos si bil ity of pre cise iden ti fi ca tion is pre cluded, nchen (1955/4/3). Compar a tively good pres er va tion of the shells be cause of the lack of adult spec i mens, but the shells from Zwie- per mitted analy sis of the band ing pat tern on the whorls, partic u- rzyniec match the de scrip tion and fig ures of A. subnitens from larly clearly vis ible in UV-light (apart from the main band located Mörsingen and some other Eu ro pean lo cal i ties (Gottschick and just above the pe riph ery, there are two more – above and below the Wenz, 1916; Gottschick, 1920; Schlickum, 1976, 1978). On the for mer) pre sented by Górka (2008a). It dif fers somewhat from the other hand, the system atic sta tus of A. subnitens re quires clar i fi ca- traces of pig ment bands, ob served by Binder (2008) on spec i mens tion, as it is con fused with some other spe cies. Lueger (1981) il le - from Mörsingen, which have only one ad di tional band be low the git i mately placed A. subnitens in syn on ymy of Aegopinella orbi- main one. cularis (= He lix orbicularis Klein, 1846), ig nor ing Klein’s re - The spe cies is known from many Mid dle Mio cene sites in Eu - marks concern ing the shell-surface striation of the latter species rope (Wenz, 1923b; Binder, 2008), but this is the first record for (Klein, 1846, p. 71) and the fact that Wenz (1919) trans ferred He - Po land. lix orbicularis Klein to Gonyodiscus (), and then to Pleurodiscus, fam. Pleurodiscidae (Wenz, 1923d). Re cently, the Ge nus Cepaea Held, 1837 spec i mens, iden ti fied by Lueger (1981) as A. orbicularis , were as- Type spe cies: He lix nemoralis Linnaeus, 1758 signed to A. reussi (Hörnes, 1856) in Harzhauser and Binder (2004) and later to A. reussi (Schlosser, 1907) in Harzhauser et al. (2011). Cepaea sylvestrina gottschicki Wenz, 1919 This is the first record of the spe cies in Po land, with excep tion Fig. 6E–G of a du bi ous re cord by WoŸny (1976) from Orawa – Nowy Targ *1919 Cepaea gottschicki sp. nov. – Wenz: p. 70. Ba sin. 1920 (1919) Cepaea sylvestrina gottschicki (Wenz) – Wenz: p. 152, figs 7–8. Family HELICIDAE Rafinesque, 1815 Ma te rial: 48 more or less damaged spec i mens (ZC C1–ZC C23). Ge nus Klikia Pilsbry, 1894 Di men sions (of the best pre served shell, ZC C12 ): H = 16.0 mm; B Type spe cies: He lix osculum Thomae, 1845 = 20.0 mm; h = 7.5 mm; b = 10 mm; HLW = 13.5 mm; NW = 4.75. Remarks: The tax o nomic po si tion of this form was un cer tain, but Klikia giengensis (Klein, 1846) fi nally Wenz (1920 (1919)) rec ognized the spec i mens from Stein- Fig. 6A heim, de scribed pre vi ously as C. gottschicki (Wenz, 1919), as a sub spe cies of C. sylvestrina (Schlotheim, 1820). The form of the *1846 He lix Giengensis sp. nov. – Klein: p. 69, pl. 1, figs 9a–b. ap er ture in C. sylvestrina is rather variable and more or less nar- 1874 He lix (Gonostoma) osculum var. Giengensis Krauss – rowed, as in the Recent C. nemoralis (Linnaeus, 1758), which is Sandberger, p. 585, pl. 29, figs 4–4b. re garded as a descen dant of the for mer species (Wenz, 1920). The 1911 Klikia giengensis Krauss – Wenz, p. 85, pl. 2, figs 6–10. band ing pat tern of the shells from Zwierzyniec was re vealed by 1976 Klikia (Klikia) giengensis (Klein) – Schlickum, p. 16, pl. view ing them in UV light (Górka, 2008a). 4, fig. 58. Ma te rial: One damaged spec i men and seven frag ments (includ ing Helicidae indet. a frag ment of body whorl with an ap er ture and a thick ened peri - Apart from the species recog nized here, some unidentificable frag- stome pre served) (Ml/1070). ments of other shells were found. Most fre quently, they are frag - Di men sions: H = ca. 4.5 mm; B = ca. 8.78 mm; h = ca. 3.27 mm; b ments of the up per part of the spire or basal parts with the umbi- = ca. 4.43 mm; HLW = ca. 4.22 mm; NW = 5. licus and a fragment of the ap er ture. Part of them proba bly be longs De scrip tion: The shell is flat tened with a rather broadly open um- to Protodrepanostoma sp. or to Helicodonta sp. Other may rep re- bi li cus, and a reflexed ap er ture edge with a thick peri stome. The sent a species of Leucochroopsis. whorls are sep a rated by a deeply in cised su ture and cov ered by very fine, dense papillae, also vis i ble on the base. Re marks: The sin gle spec i men from Zwierzyniec does not dif fer from those pre sented by Sandberger (1874), Wenz (1911) and DIS CUS SION Schlickum (1976). Palaeoenvironmental im pli ca tions

Ge nus Pseudochloritis C. Boettger, 1909 The eco log i cal re quire ments of ex tant mem bers of the Type spe cies: He lix incrassatus Klein, 1853 genera, which are ex tinct either in Europe or in Central Eu- rope, make it possi ble to draw in ferences about the envi ron - Pseudochloritis incrassata (Klein, 1853) ment that ex isted in the cen tral-south ern Poland of the pres- Fig. 6B–D ent day dur ing the Mid dle Mio cene. At Zwierzyniec, ter res - trial gastro pods were found together with freshwa ter spe- *1853 He lix incrassata sp. nov. – Klein: p. 208, pl. 5, fig. 6. cies, in di cat ing a high de gree of het er o ge ne ity of the lo cal- 1916 Tropidomphalus incrassatus (Klein) – Gottschick and ity. The gastro pod shells were washed down from the coast, Wenz, p. 55. proba bly by heavy rains that formed rapid torrents, dis - 2008 Pseudochloritis incrassata (Klein) – Binder, p. 172. charging to the sea. The gas tropod-bear ing sed iments might Ma te rial: Two well-pre served spec i mens and over 30 frag ments be regarded as depos its of the un derwa ter part of a delta, (ZC H1-ZC H4) and the gas tro pod as sem blage may re flect hab i tats, sit u ated Di men sions: H = 14–16 mm; B = 24–26 mm; h = 9.5–12.0 mm; b = 14.0–17.5 mm; HLW = 12.5–15.0 mm; NW = 4.75–5. both near and far away from the place of depo si tion. Re marks: The shell form and size, the appear ance of the umbi li - The fact that the assem blage contains hygrophilous and cus and the surface sculpture of the speci mens from Zwierzyniec xerophilous species, as well as species of shady and open match those of the P. incrassata shells from Mörsingen, stored in places, dem on strates that they come from vari ous habi tats. the Paläontologisches Mu seum of Bayrische Staatssammlung Mü- Most of them rep re sent gen era which have their Re cent 196 E. STWORZEWICZ ET AL.

Fig. 6. Land gas tro pods from Zwierzyniec; scale bars 5 mm. A. Klikia giengensis (Klein, 1846), Ml/1070; B–D. Pseudochloritis incra- ssata (Klein, 1853), ZC H1; E–G. Cepaea sylvestrina gottschicki Wenz, 1919, ZC C12 MIO CENE TER RES TRIAL GAS TRO PODS FROM ZWIERZYNIEC 197

members living in shrubland and/or in forests, more or less Ac knowl edge ments hu mid, and they are not lim ited to coastal areas (Carychium, The au thors are very grate ful to Rob ert A.D. Cameron (Uni - Leiostyla, Azeca, Ver tigo, Dis cus, Helicodiscus, Hawaiia, ver sity of Shef field and Natu ral His tory Museum, Lon don, United Phenacolimax, Vitrea, Clausiliidae and some of the Helici- Kingdom) for his crit i cal reading of an ear lier ver sion of the manu - dae). Another group consists of gastro pods, which are char- script and for cor rect ing the Eng lish. The au thors also thank acter is tic of open, warm and dry grasslands, mostly on a cal- Andrea Peresvet-Soltan (In sti tute of Sys tem at ics and Evo lu tion of cium-rich sub strate, with insolated cal car e ous rocks and Ani mals, Pol ish Acad emy of Sci ences, Kraków, Poland) for pre - xerophilous veg e ta tion (Pupilla, Granaria, Pupoides, partly par ing the mi cro scope pho tos. Spe cial thanks are due three re view - Vallonia). The mod ern dis tri bu tion of Gastrocopta and ers Beata M. Pokryszko (Museum of Natu ral His tory, Wroclaw Strobilops overlaps with the zone of subtrop i cal mixed me- Uni ver sity), Tadeusz Peryt (Pol ish Geo log i cal In sti tute, Na tional sophytic and swamp forests, es sentially ever green, but with Research Insti tute) and Mathias Harzhauser (Natu ral His tory Mu- seum, Geo log i cal-Paleontological De part ment, Vi enna) for many dis tinctly marked sea sonal changes of temper ature. valu able com ments and sug ges tions. In the Zwierzyniec gas tro pod as semblage, there are also members of genera that are repre sented in mod ern fau nas only by rare species with poorly known ecolog i cal require - ments. For ex ample, sev eral Microstele spe cies are re - REF ER ENCES stricted to the arid ar eas of trop ical Af rica, In dia and Sri Lanka (for merly Cey lon), but their eco log i cal re quire ments Al der, J. A., 1830. Cat a logue of the land and fresh wa ter testaceous Mollusca found in the vi cin ity of Newcas tle-upon-Tyne. Tra- seem to be known only for a species from Ceylon, which nsactions of the Nat ural History So ci ety of Northumberland, was found “in old posts and on palmyra trees” (Pilsbry, 1: 26–41. 1920–21). Four recent species of Negulus are known only Alexandrowicz, S. W., Garlicki, A. & Rutkowski, J., 1982. Pod- from the moun tain areas of tropi cal Africa and they were stawowe jednostki litostratygraficzne miocenu zapadliska found in the leaf lit ter of ever green forests, under bark, and przedkarpackiego. Kwartalnik Geologiczny, 26: 470–471. [In in bushes, between 800 and 4000 m a.s.l. (Bruggen, 1994). Pol ish]. The only spe cies, which unequiv o cally indi cates a close Andreae, A., 1902. Zweiter Beitrag zur Binnenconchylienfauna proxim ity to the sea, is the uniden ti fied species of Myoso- des Miocäns von Oppeln in Schlesien. Mitteilungen aus dem tella. The Recent repre sen ta tives of this genus live on litto - Roemer-Mu seum, 18: 1–31. ral rocks and stones, and in detri tus, mainly above the high- Andreae, A., 1904. Dritter Beitrag zur Kenntnis des Miocäns von tide mark (Martins, 1996). The other ellobiids, found in Oppeln i. Schl. Mitteilungen aus dem Roemer-Mu seum, 20: 1–22. Zwierzyniec, belong to Carychium, a group of terres trial Benson, W. H., 1856. New land shells col lected by E. L. Layard, snails with a prefer ence for humid habi tats, living both in Esq., and de scribed by W. H. Benson, Esq. An nals and Mag a - forests and in meadows. zine of Nat u ral History , Se ries 2, 18: 433–439. The fresh wa ter gas tro pods are rep re sented mainly by Binder, H., 2002. Die land- und Süsswassergastropoden aus dem lymnaeids of the genus Galba, which in di cate the pres ence Karpatium des Korneuburger Beckens (Niederösterreich; of rela tively small and very small water bodies (Prisyazh- Untermiozän). Beiträge zur Paläontologie, 27: 161–203. nyuk et al., 2006). Binder, H., 2008. The system atic po sitions of the gen era Psudo- chloritis C. Boettger 1909 and Joosia Pfeffer 1929. Archiv für Molluskenkunde, 137: 167–193. Binney, A., 1840. A monograph of the he li ces in hab it ing the Remarks on biogeography United States. Boston Jour nal of Nat u ral His tory, 3: 421–438. Most gen era rep re sented at Zwierzyniec have Re cent Boettger, O., 1874. Ueber die Gliederung der Cyrenenmergel- gruppe im Mainzer Becken. Bericht über die Senckenbergi- rep re sen ta tives in south ern Eu rope or in dis tant, ex tra-Eu ro- sche Naturforschende Gesellschaft in Frank furt am Main, pean regions. Pomatias, Myosotella, Azeca, Leiostyla and 1873/74: 50–102. Triloba are re cently more or less widely dis tributed in the Boettger, O., 1877. Ueber die Fauna der Corbicula-Schichten im Medi ter ra nean region and in the Cauca sus. The other group Mainzer Becken. Palaeontographica, 24: 185–219. consists of gen era, which have living mem bers distrib uted Boettger, O., 1884. Fossile Binnenschnecken aus den untermio- nearly world-wide with the excep tion of Europe (Gas tro- cänen Corbicula-Thonen von Niederrad bei Frank furt. Be- copta), or are re stricted to North Amer ica and south-eastern richt über die Senckenbergische Naturforschende Gesell- Asia (Strobilops, Hawaiia), to trop ical Africa, India and schaft in Frank furt am Main, 1884/1885: 258–280. Cey lon (Microstele), or only to tropi cal Africa (Negulus). Boettger, O., 1886 (1885/1886). Beiträge zur Herpetologie und Both groups may be regarded as rem nants of a malacofauna Malakozoologie Südwest-Africas. Bericht über die Sencken- that had a wider range during the Tertiary, and one that ex- bergische Naturforschende Gesellschaft in Frank furt am isted along the northern coasts of the Tethys (Stworzewicz, Main, 1885/1886: 3–29. Boettger, O., 1889. Die Entwicklung der Pupa-Arten des Mittel- 1993). rheingebietes in Zeit und Raum. Jahrbücher des Nassau- The most numer ous group contains the members of ge- ischen Vereins für Naturkunde, 42: 225–327. nera, which have extant conge ners in Poland (Carychium, Boettger, O., 1908. Die fossilen Mollusken der Hydrobienkalke Pupilla, Granaria, Vallonia, Vertigo, Discus, Vitrea, Aego- von Budenheim bei Mainz. Nachrichtsblatt der Deutschen pinella and Cepaea). Only a few genera are to tally extinct Malakozoologischen Gesellschaft, 40: 145–157. (Serrulastra, Miozonites, Pseudochloritis) and are thus less Bouchet, P. & Rocroi, J. P., 2005. Classication and nomenclator of use ful for biogeographical con sid er ations. gas tro pod fam i lies. Malacologia, 47: 1–397. 198 E. STWORZEWICZ ET AL.

Bourguignat, J. R., 1881. Histoire Malacologique de la Colline de lusca: Gastropoda: Vallonidae). Schriften zur Malakozoolo- Sansan. G. Masson, Paris, 175 pp. gie, 8: 1–227. Bruggen, A. C., 1994. Revisionary notes on Negulus O. Boettger, Gottschick, F., 1911. Aus dem Tertiärbeckens von Steinheim a. A. 1889, a ge nus of minute Af ri can land snails (Gastropoda Jahreshefte des Vereins für Vaterländische Naturkunde in : Vertiginidae). Zoologische Mededelingen, 68: 5– Württemberg, 67: 496–534. 20. Gottschick, F., 1920. Die land- und Süsswassermollusken des Clessin, S., 1877. Die tertiären Binnenconchylien von Undorf. Tertiärbeckens von Steinheim am Albuch. Archiv für Mollus- Correspondenz-blatt des Zoologisch-mineralogischen Ve- kenkunde, 52: 33–47. reins zu Regensburg, 31: 34–41. Gottschick, F. & Wenz, W., 1916. Sylvanaschichten von Hohen- Czapowski, G., 1984. Bar rier rocks in Up per Mio cene at southern memmingen und ihre Fauna. Nachrichtsblatt der Deutschen margin of the Holy Cross Mts. Przegl¹d Geologiczny, 32: Malakozoologischen Geselschaft, 48: 17–31, 55–74, 97–122. 185–194. [In Pol ish, Eng lish summary]. Gottschick, F. & Wenz, W., 1919. Die Land- und Süsswasser- Czapowski, G. & Studencka, B., 1990. The sedimentological- mollusken des Tertiärbeckens von Steinheim am Aalbuch. I. palaeontological study of the Lower Sarmatian bar rier de pos- Die Vertiginiden. Nachrichtsblatt der Deutschen Malakozoo- its near Chmielnik (south ern bor der of the Holy Cross Mts). logischen Geselschaft, 51: 1–23. Przegl¹d Geologiczny, 38: 117–127. [In Pol ish, Eng lish sum- Gozhik, P. F. & Prisyazhnyuk, V. A., 1978. Presnowodnye i na- mary]. zemnye molluski miocena prawoberežnoj Ukrainy. Naukova Draparnaud, J. P. R., 1801. Tableau des Mollusques Terrestres et Dumka, Kiev, 174 pp. Fluviatiles de la France. Renaud – Montpellier, Bossange, Górka, M., 2003. Rozwój facjalny tzw. sarmatu detrytycznego na Masson et Besson – Paris, 116 pp. obszarze wschodniego Ponidzia. Unpub lished Ph.D. The sis, Dudziak, J. & £aptaœ, A., 1991. Stratigraphic po sition of Mio cene In sti tute of Ge ol ogy, Uni ver sity of War saw, Po land, 100 ms. car bon ate-siliciclastic de pos its near Chmielnik (Œwiêto- pp. [In Pol ish]. krzyskie Mountains area, Cen tral Po land) based on calca- Górka, M., 2008a. Shell colour pat tern in two fossi helicid snails, reous nannofossils. Bulle tin of the Polish Academy of Earth Tropidomphalus incrassatus (Klein, 1853) and Cepaea syl- Sci ences, 39: 55–66. vestrina gottschicki Wenz, 1919, from the Mid dle Mio cene of Dupuy, D., 1850. De scrip tion de quelqes espÀces de coquilles Po land. Acta Geologica Polonica, 58: 105–111. terrestres fossiles de Sansan. Jour nal de Conchyliologie, 1: Górka, M., 2008b. “Sarmat detrytyczny” wschodniego Ponidzia: 300–315. litofacje i œrodowiska sedymentacji. In: Wojewoda, J. (ed.), Eichwald, E., 1829. Zoologia Specialis. Josephi Zawadzki, Vilnae, Baseny œródgórskie. Kontekst regionalny œrodowisk i proce- 334 pp . sów sedymentacji. Polska Konferencja Sedymentologiczna, Eichwald, E., 1830. Naturhistorische Skizze von Lithauen, Volhy- Kudowa-Zdrój, 15-21 wrzeœnia 2008. WIND, Wroc³aw, pp. nien und Podolien in geognostisch-mineralogischer, botani- 5–6. [In Pol ish]. scher und zoologisher Hinsicht. Josephi Zawadzki, Vilnae, Grateloup, M., 1827. Tab leau des co quilles fos sils qu’on rencontre 256 pp. dans les terrains calcaires tertiaires (faluns) des envi rons de Eichwald, E., 1853. Lethaea Rossica ou Paléontologie de la Ru- Dax, dans le département des Landes. Bul le tin d’Histoire ssie, DerniÀre Période. E. Schweizerbart, Stuttgart, 518 pp. Naturella de la Société Linnéenne, 2: 72–109. Férussac, A. E. J.d’Audebard de, 1807. Essai d’une Méthode Harzhauser, M. & Binder, H., 2004. Synop sis of the Late Miocene Conchyliologique Appliqueé aux Mollusques Fluviatiles et mollusc fauna of the clas sical sec tions Richardhof and Eich- Terrestres d’Aprés la Cosidération de l’ et de son Test. kogel in the Vienna Ba sin. Archiv für Molluskenkunde, 133: Delance, Paris, 162 pp. 109–165. Férussac, A. E. J.d’Audebard., de, 1821–1822. Ta bleaux Systé- Harzhauser, M., Gross, M. & Binder H., 2008. Biostratigraphy of matiques des Animaux Mollusques Classés en Familles Natu- Middle Miocene (Sarmatian) wetland systems in an Eastern relles, dans Lesquels on a êtabli la Con cor dance de tTous les Al pine intramontane ba sin (Gratkorn Ba sin, Austria): the ter - SystÀmes; Suivis d’un Prodome Générale Pour Tous les Mol- res trial gas tro pod ap proach. Geologica Carpathica, 59: 45– 58. lusques, Terrestres ou Fluviatiles, Vivants et Fossiles. Ar- Harzhauser, M., Daxner-Hock, G., Göhlich, U. B. & Nagel, D. thus-Bertrand, Paris, 114 pp. 2011. Complex fau nal mix ing in the early Pannonian palaeo- Finger, I., 1998. Gastropoden der kleini-Schichten des Steinhei- Danube Delta (Late Mio cene, Gaweinstal, Lower Austria). mer Beckens (Miozän, Süddeutschland). Stuttgarter Beiträge Annalen des Naturhistorischen Muse ums in Wien, Serie A, zur Naturkunde Ser. B, 259: 1–51. 113: 167–208. Fischer, K., 1920. Ein neuer Pupoides aus den obermiocänen Hörnes, M., 1856. Die fossilen Mollusken des Tertiär-Beckens Landschneckenmergein von Frank furt a. M. Archiv für Mol- von Wien. I. Univalven. Abhandlungen der Geologischen luskenkunde, 52: 92–94. Reichsanstalt, 3: 461–736. Fischer K. & Wenz, W., 1914. Die Landschneckenkalke des Main- Jickeli, C. F., 1874. Fauna der Land- und Süsswasser-Mollusken zer Beckens und ihre Fauna. Jahrbücher des Nassauischen Nord-Ost-Afrika’s. Nova Acta der Kaiserlich Leopoldinisch- Vereins für Naturkunde, 67: 22–154. Carolinisch Deutschen Akademie der Naturforscher, 37: 1– Friedberg, W. S., 1905. Eine sarmatische Fauna aus der Umge- 352. gend von Tarnobrzeg in Westgalizien. Sitzungsberichten der Klein, A., von, 1846. Conchylien der Süsswasserkalkformationen kaiserlichen Akademie der Wissenschaften in Wien, Mathe- Württembergs. Jahreshefte des Vereins für vaterländische matisch-naturwissenschaftliche Klasse, 114: 275–327. Naturkunde in Württemberg, 2: 60–116. Friedberg, W. S., 1928. Études sur le miocÀne de la Pologne – Klein, A., von, 1853. Conchylien der Süsswasserkalkformation partie IV. Kosmos, 53: 69–81. [In Pol ish, French summary]. Württembergs. Jahreshefte des Vereins für vaterländische Gaál, S., 1911. Die sarmatische Gastropodenfauna von Rákosd im Naturkunde in Württemberg, 9: 203–223. Komitat Hunyad. Mitteilungen aus dem Jahrbuch der köni- Kókay, J., 2006. Nonmarine mollusc fauna from the Lower and glichen ungarischen geologischen Reichsanstalt, 18: 1–112. Middle Mio cene, Bakony Mts, W Hungary. Geologica Hun- Gerber, J., 1996. Re vi sion der Gattung Vallonia Risso 1826 (Mol- garica, Se ries Palaeontologica, 56: 1–196. MIO CENE TER RES TRIAL GAS TRO PODS FROM ZWIERZYNIEC 199

Küster, H. C., 1887. Die Schliessschnecken und die verwandten Morton, J. E., 1955. The evo lu tion of the Ellobiidae with a dis cus - Gattungen (Clausilia, Balea, Cylindrella, Megaspira) in Ab- sion on the or i gin of the Pulmonata. Pro ceed ings of the Zoo - bildungen nach der Natur mit Beschreibungen. Systemati- log i cal So ci ety of Lon don, 125: 127–168. sches Conchylien-Cabi net von Mar tini und Chemnitz, 1: 1– Müller, O. F., 1774. Vermium Terrestrium et Fluviatilium, seu 355. Animalium Infusoriorum, Helminthicorum, et Testaceorum, Lindholm, W. A., 1913. Beschreibung neuer Arten und Formen non Marinorum succincta Historia. Heineck et Faber, Hav- aus den Kaukasus-Gebiete. Nachrichtsblatt der Deutschen niae et Lipsiae, 214 pp. Malakozoologischen Geselschaft, 45: 17–23, 62–69. Nordsieck, H., 1981. Fossile Clausilien, V. Neue Taxa neogener Lindholm, W. A., 1926. Ein Beitrag zur Kenntnis der Mollus- europäischer Clausilien, II. Archiv für Molluskenkunde, 111: kenfauna der Krim. Archiv für Molluskenkunde, 58: 161–177. 63–95. Linnaeus, C., 1758. Systema Naturae, Vol ume 1. Laurentii Salvii, Nordsieck, H., 2007. Worldwide Door Snails. ConchBooks, Hac- Holmiae, 823 pp. kenheim, 214 pp. Lowe, R. T., 1852. Brief di ag nostic no tices of new Maderan land Pfeif fer, L., 1857. Neue Landschnecken. Malakozoologische Blät- shells. The An nals and Maga zine of Nat ural History , 9: 275– ter, 4: 85–89. 279. Pilsbry, H. A. 1920–1921. Pupillidae (Vertigininae, Pupillinae. In: Ložek, V., 1964. Neue Mollusken aus dem Altpleistozän Mittel- Pilsbry, H. A. (ed.), Man ual of Con chol ogy, Vol ume 26. Con- europas. Archiv für Molluskenkunde, 93: 193–199. chological Depart ment, Academy of Natu ral Sciences of Phil- Lueger, J. P., 1981. Die Landschnecken im Pannon und Pont des a del phia, Phil a del phia, 254 pp. Wiener Beckens. Österreichische Akademie der Wissenschaf- Prisyazhnyuk, V. A., 1971. Molluski rodu Microstele O. Boettger ten, mathematisch-naturwissenschaftliche Klasse, Denkschrif- (Pulmonata). Dopovidi Akademii Nauk Ukrainskoi RSR, Se- ten, 120: 1–124. riya B, 7: 599–601. Luleva, S. A. & Prisyazhnyuk, V. A., 1990. Novye dannye po Prisyazhnyuk, V. A., 1972. Nekotoriye predstaviteli nazemnich paleontologicheskoj harakteristike miocenowyh otlozhenij w molluskov v sarmatzkich otlozhenyach Volyno-Podoli. Geo- rajone r. Buglovki (zapadnye rajony USSR). Doklady Aka- logicheskiy Zhurnal, 32: 130–133. [In Ukrai nian]. demii Nauk Ukrainskoy SSR, Seriya B, 7: 34–38. [In Rus sian, Prisyazhnyuk, V. A., 2008a. New lo cal i ties of ter res trial mol lusks Eng lish sum mary]. in the Mio cene de posits of the Eastern Dnie per Ba sin, south - £omnicki, A. M., 1886. S³odkowodny utwór trzeciorzêdny na ern Ukraine. Biostratigraphichni osnovi pobudovi stratigra- Podolu galicyjskim. Sprawozdania Komisji Fizyjograficznej, phizhnich shem phanerozoju Ukraini. Zbirnik Naukovich 20: 48–119. [In Pol ish]. Pratz Institutu Geologizhnich Nauk NAN Ukraini, Natsio- £uczkowska, E. & Rutkowski, J., 1970. Pozycja stratygraficzna nalna Akademia Nauk Ukraini, Paleontologichne Towaris- detrytycznych osadów sarmatu na po³udniowym obrze¿eniu tvo, Kiiw, pp. 230–243. [In Ukrai nian]. Gór Œwiêtokrzyskich. Sprawozdania z Posiedzeñ Komisji Prisyazhnyuk, V. A., 2008b. Ter res trial and fresh wa ter molluscs Naukowych Polskiej Akademii Nauk, Oddzia³ w Krakowie, of Trijebine, Sjenica (south-west ern Ser bia). Bul le tin de 13: 563–566. [In Pol ish]. l’Académie Serbe des Sciences et des Arts, Classe des Sci- Manganelli, G. & Giusti, F., 2000. The gastrocoptids of the Fossil ences mathématiques et naturelles, Sci ences naturelles, 44: Forest of Dunarobba (Cen tral It aly) and a pre limi nary re vi - 85–109. sion of the Eu ro pean Ter tiary nomi nal spe cies of Albinula and Prisyazhnyuk, V. A., Kovalenko V. A, Górka M. & Stworzewicz Vertigopsis (Gastropoda Pulmonata: Gastrocoptidae). Bolle- E., 2006. The fresh-wa ter gas tro pods from the Sarmatian tino della Societ´ Paleontologica Italiana, 39: 55–82. (Lymnaeidae, Bulinidae, Planorbiidae) of Zwierzyniec (Cen- Manganelli, G., Aldinucci, M., Capezzuoli, E. & Benocci, A., tral Po land). Problemy paleontologii ta biostratigraphii prote- 2008 (2007). Strobilopsid land snails (Gastropoda: Pulmo- rozoju i phanerozoju Ukraini. Zbirnik Naukowich Pratz Insti- nata, Strobilopsidae) from the late Messinian – Early Pleis to- tutu Geologizhnich Nauk NAN Ukraini, Natsionalna Akade- cene (?) of Balze di Caspreno (central Italy). Bolletino della mia Nauk Ukraini, Paleontologichne Tovaristwo, Kiiw, pp. Societ´ Paleontologica Italiana, 46: 101–106. 254– 264. [In Ukrai nian]. Mar tins, A. M. F., de 1996. Anat omy and sys temat ics of the West - Reuss, A. E., 1849. Geognostische Skizze der tertiären Süsswas- ern Atlan tic Ellobiidae (Gastropoda: Pulmonata). Malaco- serschichten des nördlichen Böhmens. Palaeontographica, 2: logia, 37: 163–332. 1–42. Matheron, M. P., 1832. Ob ser va tions sur les ter rains tertiares du Reuss, A. E., 1860. Die fossilen Mollusken der tertiären Süss- Departement des Bouches-du-RhÛne et de scrip tion des Co - wasserkalke Böhmens. Sitzungsberichte der kaiserische Aka- quilles fossiles inédites ou peu com munes qu’ils renterment. demie der Wissenschaften in Wien, 42: 55–85. Annales des Sci ences et de l’Industries du Midi de la France, Riedel, A., 1963. Ein rezenter Hawaiia-Fund aus Af ghan i stan und 3: 39–61. ein fossiler aus dem Kaukasus (Gastropoda, Zonitidae). Menke, C. T., 1828. Syn opsis methodica molluscorum generum Annales Zoologici, 21: 33–41. omnium et specierum earum quae in museo Menkeano adser- Rutkowski, J., 1976. De tri tal Sarmatian de posits on the south ern vantur. Uslar, Pyrmonti, 91 pp. margin of the Holy Cross Mountains (south ern Po land). Michaud, M., 1862. De scrip tion des Co quilles fossiles des en vi - Prace Geologiczne Polskiej Akademii Nauk, Oddzia³ w Kra- rons de Hauterive (DrÛme). Jour nal de Conchyliologie, 10: kowie, 100: 1–71. [In Pol ish, Eng lish summary]. 58–84. Sandberger, F., 1858–1863. Die Conchylien des Mainzer Tertiär- Miller, K., 1900. Die Schneckenfauna des Steinheimer Obermio- beckens. C. W. Kreidel’s Verlag, Wiesbaden, 468 pp. cäns. Jahreshefte des Vereins für vaterländische Naturkunde Sandberger, F., 1870–1875. Die Land- und Süsswasser-conchylien in Württemberg, 56: 385–406. der Vorwelt. C. W. Kreidel’s Verlag, Wiesbaden, 1000 pp. Montagu, G., 1803. Testacea Bri tan nica, or Nat u ral His tory of Sandberger, F., 1886. Bemerkungen über fossile Conchylien aus Brit ish Shells, Ma rine, Land, and Fresh-wa ter, In clud ing the dem Süsswasserkalke von Leobersdorf bei Wien (Inzers- Most Min ute: Sys tem at i cally Ar ranged and Em bel lished with dorfer Schichten). Verhandlungen der kaiserlich-königlichen Fig ures. J. S. Hollis, Rom sey, 645 pp. geologischen Reichstanstalt (Wien), 14: 331–332. 200 E. STWORZEWICZ ET AL.

Say, T., 1817. De scrip tions of new spe cies of land and fresh wa ter (Gastropoda Pulmonata: Endodontodae). Ge nus, 8: 197–204. shells of the United States. Jour nal of the Academy of Nat ural Stworzewicz, E. & So³tys, Z., 1996. Miocene land snails from Sci ences of Phil a del phia, 1: 123–126. Be³chatów (Cen tral Po land). II. Aciculidae (Gastropoda Schileyko, A. A., 1984. Nazemnye molluski podotriada Pupillina Prosobranchia). Paläontologische Zeitschrift, 70: 67–77. fauny SSSR (Gastropoda, Pulmonata, Geophila). In: Aka- Œliwiñski, M., B¹bel, M., Nejbert, K., Olszewska-Nejbert, D., demia Nauk SSSR (ed.), Fauna SSSR, Molluski 3, 3. Nauka, G¹siewicz, A., Schreiber, B. C., Benowitz, J. A. & Layer, P., Le nin grad, 399 pp. [In Rus sian]. 2012. Badenian–Sarmatian chronostratigraphy in the Pol ish Schlickum, W. R., 1976. Die in der pleistozänen Gemeinde- Carpathian Foredeep. Palaeo ge ogra phy, Palaeoclimatology, kiesgrube von Zwiefaltendorf a.d. Donau abgelagerte Mollus- Palaeo ec ol ogy, 326–328: 12–29. kenfauna der Silvanaschichten. Archiv für Molluskenkunde, Thomae, C., 1845. Fossile Conchylien aus den Tertiärschichten 107: 1–31. bei Hochheim und Wiesbaden, gesammelt und im Naturhisto- Schlickum, W. R., 1978. Zur oberpannonen Molluskenfauna von rischen Mu seum zu Wiesbaden aufgestellt. Jahrbücher des Öcs, I. Archiv für Molluskenkunde, 108: 245–261. Nassauischen Vereins für Naturkunde, 2: 127–166. Schlickum, W. R., 1979. Helicodiscus (Hebetodiscus) ein altes Vest, W., 1867. Ueber den Schliessapparat der Clausilien. Ver- europäischen Faunenelement. Archiv für Molluskenkunde, handlungen und Mittheilungen des siebenbürgischen Vereins 110: 67–70. für Naturwissenschaften, 18: 5–18, 161–174, 188–196. Schlosser, M., 1907. Die Land- und Süsswassergastropoden vom Wenz, W., 1911. Gonostoma (Klikia) osculum Thom. und ihre Eichcogel bei Mödling. Jahrbuch der kaiserlich-königlichen Verwandten im mitteleuropäischen Tertiär. Jahrbücher des geologischen Reichstanstalt, 57: 753–791. Nassauischen Vereins für Naturkunde, 64: 75–101. Schlotheim, E. F. von, 1820. Die Petrefaktenkunde auf ihrem Wenz, W., 1915. Die fossilen Arten der Gattung Strobilops Pilsbry jetzigen Standpunkte durch die Beschreibung seiner Sam- und ihre Beziehungen zu den lebenden. Neues Jahrbuch für mlung versteinerter und fossiler Ueberreste des Thier- und Mineralogie, Geologie und Palaeontologie, 2: 63–88. Pflanzenreichs der Vorwelt erläutert. Becker, Gotha, 438 pp. Wenz, W., 1919. Zur Nomenklatur tertiärer Land- und Süsswas- Schnabel, T., 2012. A fos sil Triloba spe cies from Mac e do nia and sergastropoden. Nachrichtsblatt der Deutschen Malakozoolo- its re la tions to the re cent spe cies of the ge nus (Gastropoda: gischen Gesellschaft, 51: 68–76. Pulmonata: Clausiliidae). Archiv für Molluskenkunde, 141: Wenz, W., 1920 (1919). Helicites sylvestrinus Schlotheim. Senc- 35–42. kenbergiana, 2: 19–22 Schütt, H., 1967. Die Landschnecken der untersarmatischen Ris- Wenz, W., 1920. Die Ellobiiden des Mainzer Beckens. Senc- soenschichten von Hollabrunn, N.-Ö. Archiv für Mollusken- kenbergiana, 2: 189–192. kunde, 96: 199–222. Wenz, W., 1921. Das Mainzer Becken und seine Randgebiete. Steklov, A. A., 1966. Nazemnye Mollyuski Neogena Predkavkazya Willy Ehrig, Hei del berg, 352 pp. i ikh Stratigraphicheskoye Znachenie. Nauka, Moskva, 262 Wenz, W., 1922. Eine neue Lauria aus dem Obermiocän von pp. [In Rus sian]. Steinheim am Albuch. Archiv für Molluskenkunde, 54: 106– Steklov, A. A. & Tsytovich, M. V., 1967. O nahodke neogenovych 109. nazemnych molluskov v dolinie Kostankol v Tsentralnom Wenz, W., 1923a. Gastropoda extramarina tertiaria. In: Diener, C. Kasakhstanie. Biuleten Moskovskogo Obschchestva Ispita- (ed.), Fossilium Catalogus, I. Animalia, 17. W. Junk, Berlin, teley Prirody, Otdel Geologicheskiy, 3: 108–119. [In Rus - pp. 1–352. sian]. Wenz, W., 1923b. Gastropoda extramarina tertiaria. In: Diener, C. Strauch, F., 1977. Die Entwicklung der europäischen Vertreter der (ed.), Fossilium Catalogus, I. Animalia, 18. W. Junk, Berlin, Gattung Carychium O. F. Müller seit dem Miozän (Mollusca: pp. 353–736. Basommatophora). Archiv für Molluskenkunde, 107: 149– 193. Wenz, W, 1923c. Gastropoda extramarina tertiaria. In: Diener, C. Studencka, B., (in press). Ma³¿e z osadów œrodkowego miocenu (ed.), Fossilium Catalogus, I. Animalia, 20. W. Junk, Berlin, zapadliska przedkarpackiego: wiercenia Busko (M³yny) pp. 737–1068. PIG-1 i Kazimierza Wielka (Donosy) PIG-1. Stratygrafia i Wenz, W., 1923d. Gastropoda extramarina tertiaria. In: Diener, C. taksonomia. Biuletyn Pañstwowego Instytutu Geologicznego. (ed.), Fossilium Catalogus, I. Animalia, 21. W. Junk, Berlin, Studer, S., 1820. Kurzes Verzeichniss der bis jetzt in unserm pp. 1069–1420. Vaterlande entdeckten Conchylien. Naturwissenschaftlicher Wenz, W., 1927. Weitere Beiträge zur Fauna der pontischen Anzeiger der Allgemeinen Schweizerischen Gesellschaft für Schichten von Leobersdorf. Senckenbergiana, 9: 41–48. die Gesammten Naturwissenschaften, 3: 83–94. Wenz, W. & Edlauer, A., 1942. Die Molluskenfauna der ober- Stworzewicz, E., 1993. Palaeobiogeographical charac teris tics of pontischen Süsswassermergel von Eichkogel. Archiv für Mol- Miocene land snail fauna of Po land. Scripta Geologica, Spe - luskenkunde, 74: 82–98. cial Issue , 2: 397–406. WoŸny, E., 1976. Stra tig ra phy of the youn ger Ter tiary in the Stworzewicz, E., 1995. Miocene land snails from Be³chatów (Cen - Orawa – Nowy Targ Ba sin on the basis of fresh-wa ter and tral Po land). I. Cyclophoridae, Pomatiasidae (Gastropoda continental macrofauna. Kwartalnik Geologiczny, 20: 589– Prosobranchia). Paläontologische Zeitschrift, 69: 17–28. 595. [In Pol ish, Eng lish summary]. Stworzewicz, E., 1999a. Mio cene land snails from Be³chatów Zastawniak, E., 1980. Sarmatian leaf flora from the south ern mar - (Cen tral Po land), III. Carychiinae (Gastropoda; Pulmonata: gin of the Holy Cross Mts. (South Poland). Prace Muzeum Ellobiidae). Paläontologische Zeitschrift, 73: 261–276. Ziemi, 33: 39–107. Stworzewicz, E., 1999b. Mio cene land snails from Be³chatów Zieten, C. H., 1830. Die Versteinerungen Württembergs. Verlag & (Cen tral Po land). IV: (Gastropoda Pulmonata). Lithographie der Ex pe di tion des Werkes unsere Zeit, Stut- Sys tem atic, biostratigraphic and palaeo eco logi cal stud ies. tgart, 269 pp. Folia Malacologica, 7: 133–170. Zilch, A., 1959–1960. . In: Wenz, W. (ed.), Gas tro- Stworzewicz, E. & Prisyazhnyuk, V. A., 1997. Helicodiscus (Heli- poda, 2. Gebrüder Borntraeger, Berlin-Nikolassee, pp. 1–833. codiscus) depressus (Eichwald, 1830) or Helicodiscus (Heli- codiscus) parallelus (Say, 1821) in the Neo gene of Eu rope?