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Gastropoda, Pulmonata, Gastrodontidae) from the Zanclean (Early Pliocene) of Tuscany (Central Italy

Gastropoda, Pulmonata, Gastrodontidae) from the Zanclean (Early Pliocene) of Tuscany (Central Italy

Bollettino della Società Paleontologica Italiana, 50 (3), 2011, 165-173. Modena, 30 dicembre 2011165

A new (, , ) from the Zanclean (early Pliocene) of Tuscany (central Italy)

Giuseppe Manganelli, Ivan Martini & Andrea Benocci

G. Manganelli, Dipartimento di Scienze Ambientali, Università di Siena, Via Mattioli 4, I-53100 Siena; [email protected] I. Martini, Dipartimento di Scienze della Terra, Università di Siena, Via Laterina 8, I-53100 Siena; [email protected] A. Benocci, Dipartimento di Scienze Ambientali, Università di Siena, Via Mattioli 4, I-53100 Siena; [email protected]

KEY WORDS - Gastropods, Pliocene, Zanclean, Tuscany, Janulus spadinii n. sp.

ABSTRACT - Janulus spadinii n. sp. (Gastropoda, Pulmonata, Gastrodontidae) is described herein from the Zanclean (early Pliocene) of Balze di Caspreno (Tuscany). It belongs to a Macaronesian occasionally reported from the Tertiary of Europe. The new species is very similar to the Recent Madeiran Janulus stephanophorus by virtue of its shape, dorsal sculpture and radial rows of palatal teeth. It is also apparently similar to some fossil European species, however, due to insufficient information regarding the latter, no well supported distinction may be proposed at present. A brief survey of all the nominal taxa of the species-group assigned to Janulus is also provided.

RIASSUNTO - [Una nuova specie di Janulus (Gastropoda, Pulmonata, Gastrodontidae) dal Pliocene inferiore della Toscana (Italia centrale)] - Janulus spadinii n. sp. è descritto sulla base di materiale raccolti nelle argille continentali del Pliocene inferiore delle Balze di Caspreno (Toscana). La nuova specie appartiene ad un genere attualmente esclusivo della Macaronesia, ma conosciuto per il Terziario europeo. Janulus spadinii n. sp. è molto simile a Janulus stephanophorus, una specie vivente a Madeira (le due specie condividono: forma, scultura dorsale e file radiali di denti palatali) ma ricorda anche alcune entità descritte per il Terziario europeo. Una differenziazione adeguata rispetto a questi taxa è al momento impossibile, mancando del tutto informazioni sulla struttura dei loro denti palatali. Conclude il lavoro l’elenco di tutti i taxa nominali assegnati a Janulus, inclusivo di una breve discussione sul loro status.

INTRODUCTION lacking. Moreover, this duct may not be homologous: it is absent in Pilsbryna and connects penial dart-sac and duct The genus Janulus Lowe, 1852 is regarded as a of bursa copulatrix in Gastrodonta, penis and free oviduct European palaeoendemite of the Tertiary warm climatic in Janulus, and penis and duct of bursa copulatrix in all the phases, associated with moist forest biotopes (Esu, 1999). others (however, in , a second accessory duct It survives in the Macaronesian refuges (Walden, 1983, joins duct of bursa copulatrix and free oviduct according 1984; Cameron et al., 2007) and includes only three Recent to Pilsbry, 1946; Riedel, 1980; Gittenberger et al., 1984). species: J. bifrons (Lowe, 1831) and J. stephanophorus Consequently the monophyly of the family, as currently (Deshayes, 1850) from Madeira and the probably recently conceived, is not certain, like the number of genera/ extinct J. pompilius (Shuttleworth, 1852) from La Palma, subgenera it includes (see different accounts by Hausdorf, Canary Islands (Riedel, 1980; Fontaine et al., 2007). 1998; Riedel, 1998; Schileyko, 2003; Slapcinsky & Cole, Another Madeiran species, J. calathoides (Lowe, 1863), 2004). tentatively added by Riedel (1998), is actually a species Recent Janulus species have a discoidal to trochiform of Fitzinger, 1833 (Bank et al., 2002). However, shell, angled or shouldered at periphery, with small there is no definitive evidence that the three species really , narrow ribbing on dorsal surface of teleoconch belong to the same genus, because only one of them (J. whorls and peristome simple and not reflexed. One bifrons) is known anatomically. Janulus has a holopod species (J. bifrons) is larger, with less tightly coiled and foot and distal genitalia with a penial dart-sac and an less narrowly ribbed whorls and peristome internally accessory duct connecting penis and free oviduct (Pilsbry, thickened (it strongly recalls the shells of hygromiids 1947). Holopod with apertural palatal teeth, a such as Chemnitz, 1786, Gude penial dart-sac and an accessory duct connecting penial & Woodward, 1921, etc.). Finally, at least one species (J. complex and female genitalia (duct of bursa copulatrix or stephanophorus) has radial rows of palatal teeth at 120 free oviduct) are assigned to the gastrodontids, a mainly degrees to each other, each row consisting of three-four Nearctic family including a few other genera besides teeth: one adaxial, very small and knob-like, one medial, Janulus [Gastrodonta Albers, 1850, Nastia Riedel, 1989, short and pleat-like and one abaxial, larger and inverted Poecilozonites Boettger, 1884, Pilsbryna Baker, 1929, comma-like or sometimes V-like, split into a longer outer Morse, 1864 and Zonitoides Lehmann, 1862 branch and a shorter inner branch (Pl. 1, fig. 9). (Riedel, 1998; Slapcinsky & Cole, 2004)]. Although Although the fossil record is much poorer than it the accessory duct joining penial complex and female was formerly thought (see for example Wenz, 1923), genitalia has been regarded as a unique autapomorphy sure reports spanning from the (e.g. Janulus supporting the monophyly of the gastrodontids (Hausdorf, schottleri Wenz, 1922) to the end of the Pliocene and 1998), a robust phylogeny of these land snails is still ecology of Recent species (Seddon, 2008) are in line with

ISSN 0375-7633 doi:10.4435/BSPI.2011.15 166 Bollettino della Società Paleontologica Italiana, 50 (3), 2011

Fig. 1 - Location map of the study area. a) Structural setting of southern Tuscany. b) Geological sketch of the Montaperti-Pianella area. Numbers (1-6) indicate sites sampled during the 1990s. Most of these outcrops are now covered by recent alluvial deposits, vegetation or dumped rubble.

the palaeoecological meaning attributed to the genus in part of an elongated depression (Fig. 1a) extending for the literature. about 200 km in a NNW-SSE direction. The tectonic In this paper we describe a new Janulus species from setting responsible for the origin and evolution of this the Zanclean of Balze di Caspreno (Tuscany), already basin is still debated, and has been explained in different reported as Janulus suttonensis (Wood, 1872) by De ways: according to many authors (Costantini et al., 1982; Stefani (1880). We also shortly survey all the nominal Martini & Sagri, 1993; Pascucci et al., 1999; Carmignani taxa of the species-group assigned to this genus, many of et al., 2001; Brogi, 2011) its origin is connected with an which may belong to other groups of land snails. extensional regime acting until the late Miocene, while other researchers postulate a dominantly compressional tectonic setting (Boccaletti & Sani, 1998; Finetti et al., GEOLOGICAL SETTING 2001; Bonini & Sani, 2002). Regardless of its structural origin, the Siena Basin The Siena Basin is a post-collisional basin of the inner was interested by a continental and marine deposition Northern Apennines, a collisional belt formed during during the Neogene, and to limited and discontinuous the Cenozoic in response to the interaction between the sedimentation of alluvial deposits (Aldinucci et al., 2007) Adria and the Corso-Sardinian microplates (Carmignani and travertine (Brogi et al., 2010) during the Quaternary. et al., 2001). It corresponds to approximately the middle Its Neogene basin-fill history dates back to the latest G. Manganelli et alii - A new species of Janulus from Zanclean of Tuscany 167

Miocene (Messinian) with the deposition of a thick Fm. exposed in the northern Casino Basin (Bossio et fluvio-lacustrine succession that unconformably overlies al., 1998). However, micropalaeontological analysis pre-Neogene bedrock. Most of these Miocene deposits carried out in fine floodplain sediments (Bambini, pers. are buried beneath Pliocene sediments and crop out in comm.) shows a lack of recycled Serravallian calcareous limited western marginal areas of the basin (Lazzarotto nannofossils, while bedrock-derived nannofossils are & Sandrelli, 1977; Bossio et al., 2000). Marine conditions common. These findings suggest that the deposits date in the Siena Basin began at the base of the Zanclean back to the Pliocene. According to Martini et al. (2011), (Costantini et al., 1982; Bossio et al., 1993), with the integrated palaeontological and stratigraphical analysis deposition of a thick sedimentary succession, ranging of the continental and marine succession exposed in the from nearshore (sands and conglomerates) to offshore surrounding area also enables assignment of these deposits (mud) environments (Costantini et al., 1982; Manganelli to the MPl3 Zone of planktonic foraminifera zonation et al., 2010; Martini et al., 2011). Marine sedimentation (zonal scheme of Foresi et al., 2001). persisted until the late Piacenzian, when uplifting of southern Tuscany caused emergence of the Siena Basin (Costantini et al., 1982; Bossio et al., 1993). Local SYSTEMATIC PALAEONTOLOGY episodes of continental sedimentation have been reported within the marine deposits in the southwestern sector of Janulus spadinii n. sp. the basin (Bossio et al., 1992, 1993). (Pl. 1, figs 2-7) Janulus material was collected along the Arbia River (Montaperti-Pianella area, northern sector of Diagnosis - A species of gastrodontid Janulus with the Siena Basin), where continental deposits crop out shell very similar to that of Recent Madeiran Janulus discontinuously in small scattered outcrops (Fig. 1b), stephanophorus by virtue of its shape, dorsal sculpture and most of which are now covered by recent alluvial deposits, radial rows of palatal teeth (three internal radial rows of vegetation or rubble from quarries. The best exposures teeth at 120 degrees to each other; each row consisting of of continental deposits crop out in natural cliffs on the three teeth, the more external of which larger and inverted left bank of the Arbia river (the “Balze di Caspreno”, comma-like), but differing by virtue of its lesser size and fossiliferous sites 5 and 6 in Fig. 1b). According to the more depressed shell. Martini et al. (2011) these sediments are indicative of a generic fluvial setting, with dominantly clayey floodplain Description - Shell (Pl. 1, figs 2-7) dextral, very small silts (locally containing peat horizons) and fine sands, in size, lenticular in shape, with about six, very narrow with subordinate lens-shaped bodies of fluvial gravelly and tightly coiled whorls, separated by deep sutures; last sand. The age of these deposits has long been debated: whorl angled at periphery; umbilicus small, narrow and De Stefani (1880), who first described this fossiliferous cylindrical; semilunar, with light parietal callus site, attributed these deposits to the lower part of the and peristome not thickened or reflexed; protoconch Pliocene, on the basis of lithological and palaeontological sculpture unknown; teleoconch with many strong, regular, similarities with analogous sediments exposed close to sligthly prosocline ribs, stopping just below periphery Siena, where they are interbedded with certainly Pliocene (in one specimen: 54 ribs in first teleoconch whorl, 61 in marine strata (De Stefani, 1876). De Stefani (1880), and second, 63 in third, Pl. 1, fig. 2); internally, three radial later De Castro & Pilotti (1933), also described fossil rows of palatal teeth at 120 degrees to each other; each remains of brackish gastropods in these muddy sediments, row consisting of three teeth: one adaxial, very small implying connection with a nearby marine area. Since and knob-like, one medial, small and knob-like and one marine conditions in the Siena Basin started in the earliest abaxial, larger and inverted comma-like. Pliocene, De Stefani (1880) and De Castro & Pilotti (1933) took this fact as an indirect evidence of the Pliocene age Dimensions - The largest specimen, an incomplete of these deposits. The deposits of the Balze di Caspreno shell, is 4.2 mm wide. The holotype, a juvenile shell, is area were later attributed to a generic Neogene (Signorini, 1.4 mm high and 2.8 mm wide. 1966, 1967) and recently assigned to the Messinian (Lazzarotto et al., in press), due to strong lithological Type locality and horizon - Balze di Caspreno, along similarities with the “Argille del Casino” Fm. cropping out Arbia River (Montaperti-Pianella area, east of Siena), in the nearby Casino Basin (Lazzarotto & Sandrelli, 1977). lacustrine grey clays of Zanclean age (MP13 Zone of Partially in contrast with this attribution, Manganelli et al. planktonic foraminifera zonation of Foresi et al., 2001). (2007) reported some continental molluscs ( cf. For collecting sites, see Fig. 1. romani Wenz, 1915 and aloisii Manganelli, Delle Cave & Giusti, 1989), which in Europe have been Type material - Holotype (Pl. 1, fig. 2), a juvenile recorded only from the Zanclean. shell from site 4 (Manganelli collection, Dipartimento Finally, Martini et al. (2011) reported the discovery di Scienze Ambientali, University of Siena, no. 39588); of cobbles showing evidence of bio-erosion (e.g. about 60 paratypes consisting of 110 shell fragments borings) in gravelly fluvial facies, indicating provenance (corresponding to at least 55 specimens) from Site from recycled marine nearshore gravels. According to 2 (Manganelli collection, Dipartimento di Scienze the geological history of the Siena Basin (Costantini Ambientali, University of Siena, no. 39587); 3 shells from et al., 1982; Lazzarotto et al., in press) these cobbles Site 4 (Manganelli collection, Dipartimento di Scienze could only be derived from unknown Pliocene deposits Ambientali, University of Siena, no. 39589); 7 shell or at least from the Serravallian Ponsano Sandstone fragments (corresponding to at least 5 specimens) from 168 Bollettino della Società Paleontologica Italiana, 50 (3), 2011

Site 6 (Manganelli collection, Dipartimento di Scienze the other species, most of which may belong to completely Ambientali, University of Siena, no. 39590). different genera. For example, according to Lueger (1981), J. joossi Schlickum, 1970 is a species of Discus (), Derivation of name - The new species is named namely Discus pleuradras (Bourguignat, 1881). after our dear friend Valeriano Spadini from Lucignano Among fossil species which belong or may belong (Arezzo), passionate naturalist and scholar of fossil corals to this genus, Caspreno Janulus matches the alleged and molluscs. syntype of Patula angustiumbilicata illustrated by Ferrero Mortara et al. (1984: Pl. 54, figs 1a-b),Janulus austriacus Remarks - Shell features of the species from the Harzhauser & Binder, 2004 (Pl. 7, figs 12-15) from the late Zanclean of Balze di Caspreno are sufficient to establish Miocene of Lower , suttonensis Wood, 1872 that it really belongs to Janulus: it is very similar to the (Pl. 1, figs 2a-c) from the Pliocene of England andPatula Recent J. stephanophorus (the type species of the genus) olisipponensis Roman, 1907 (fig. 8) from the Neogene with which it shares shape, dorsal sculpture and radial of Portugal. Unfortunately, at the present state of the art, rows of palatal teeth. no well supported distinction can be proposed until more In Fossilium Catalogus, Wenz (1923, pp. 300-306) detailed descriptions of these taxa become available. As listed 12 Janulus species, one of which - Janulus striatus far as we know, it differs from Janulus suttonensis, of (Eichwald, 1830) - only tentatively assigned to this genus. which we examined a detailed drawing of the holotype by Subsequently, Schlickum (1979), Schlickum & Geissert Gordon Riley, kindly made available by Richard Preece, (1980), Yü & Zhang (1982) and Harzhauser & Binder and from Janulus austriacus by virtue of its lenticular (2004) described four other putative Janulus species shape (trochiform shape in Janulus suttonensis; discoidal (see Appendix 1). Unfortunately information about shape in Janulus austriacus). It is very similar to the shells most of them, especially on the presence and structure of “Janulus” angustiumbilicatus published by Ferrero of the internal radial rows of palatal teeth, is scarce or Mortara et al. (1984), but they are different from Sacco’s absent and this prevents sure generic assignment. Only description and figures of this taxon (see Appendix). the late Miocene J. gottschicki (Jooss, 1912) and J. Finally it differs from Janulus olisipponensis, on the basis schottleri Wenz, 1922 are known to have palatal teeth but of Roman’s (1907) figure only, by virtue of its lenticular information on the former is inadequate (see Jooss, 1912). shape and slightly prosocline ribs (discoidal shape and On the contrary the pattern of palatal teeth of J. schottleri markedly prosocline ribs in Janulus olisipponensis). is consistent with that of the Recent J. stephanophorus and consequently its inclusion in Janulus is well supported. Some other species may belong to Janulus on the basis ACKNOWLEDGEMENTS of their shell shape and dorsal sculpture, but we lack information about radial rows of palatal teeth. They are: Helen Ampt revised the English. Mikaela Bernardoni J. angustiumbilicatus (Sacco, 1886, sensu Ferrero Mortara and Florence Conti (Library of the Faculty of Sciences of the et al., 1994), J. austriacus Harzhauser & Binder, 2004, J. University of Siena, Italy) helped with bibliographic research. Elena Gavetti and Daniele Ormezzano (Museo Regionale di olisipponensis (Roman, 1907) and J. suttonensis (Wood, Scienze Naturali, Turin, Italy) supported with information on L. 1872). On the contrary nothing sure can be said about all Bellardi and F. Sacco’s type-material, Richard Preece (University

EXPLANATION OF PLATE 1

Fig. 1 - Janulus stephanophorus (Deshayes, 1850). SEM apical view of a shell from Ilha da Madeira: Ribeira da Porto Novo, A.H. Walden leg. 11.11.1972 (Göteborgs Naturhistoriska Museum 72-13.386); scale bar: 2 mm.

Figs 2-7 - Janulus spadinii n. sp. from Balze di Caspreno. 2 - Holotype (a juvenile shell) from Site 4 (Manganelli collection, Dipartimento di Scienze Ambientali, University of Siena, no. 39588); SEM apical (a) and apertural (b) views; scale bar: 2 mm. 3 - A juvenile shell from Site 4 (Manganelli collection, Dipartimento di Scienze Ambientali, University of Siena, no. 39589); SEM apical view; scale bar: 2 mm. 4-7- Shell fragments from Site 2 (Manganelli collection, Dipartimento di Scienze Ambientali, University of Siena, no. 39590); SEM views of palatum illustrating the internal radial rows of palatal teeth; scale bar: 2 mm.

Fig. 8 - Janulus bifrons (Lowe, 1831): apical (a), apertural (b) and umbilical (c) views of a shell from Ilha da Madeira: Garajau, A.H. Walden leg. 2.4.1983 (Göteborgs Naturhistoriska Museum 83-16.180); scale bar: 4 mm.

Fig. 9 - Janulus stephanophorus (Deshayes, 1850): apical (a), apertural (b) and umbilical (c) views of a shell from Ilha da Madeira: Ribeira da Porto Novo, A.H. Walden leg. 11.11.1972 (Göteborgs Naturhistoriska Museum 72-13.386); scale bar: 4 mm.

Fig. 10 - Janulus pompilius (Shuttleworth, 1852): apical (a), apertural (b) and umbilical (c) views of a shell from La Palma, Canary Islands, B.F. Blauner leg. 1851 (syntype, Naturhistorisches Museum Bern 18777). Photo by Eike Neubert (© Centre Suisse de Cartographie de la Faune, reproduced by permission); scale bar: 2 mm. G. Manganelli et alii - A new species of Janulus from Zanclean of Tuscany Pl.169 1 170 Bollettino della Società Paleontologica Italiana, 50 (3), 2011

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APPENDIX - putative fossil Janulus species

Patula (Janulus) angustiumbilicata Sacco, 1886: 190-1, Pl. 1, figs Helix gyrorbis Klein, 1846: 72, Pl. 1, figs 14a-c. 17a-b. Type locality: “… bei Oepfingen in der Nähe von Ehingen...” Type locality: “Alluvioni plioceniche […] di Fossano” (Sacco, Status: uncertain. Probably a Discus species. Sandberger (1872: 1886: 169). Pl. 21, figs 7a-7b) depicted a completely different species. Status: uncertain. According to Sacco’s description and figures, the species has a punctiform umbilicus, but Ferrero Mortara et Janulus joossi Schlickum, 1979: 410 (depicted by Schlickum, 1978: al. (1984: Pl. 54, figs 1a-c) figured a putative syntype with a Pl. 19, fig. 17). small umbilicus. The material reported by Ferrero Mortara et al. Type locality: “Oberpannon, Süsswassermargel … Öcs, (1984) consists of about ten specimens, all consistent with the one Komitat Veszprém, Ungarn.” illustrated. However, there is no evidence that they constitute the Status: uncertain. According to Lueger (1981) a junior actual type material because the only label accompanying them dates of Discus pleuradras (Bourguignat, 1881). back to the 1970s (Daniele Ormezzano, pers. comm., September 2010). Although no information is available on radial rows of palatal Janulus mörsingensis Jooss, 1918: 289-90. teeth, the specimen illustrated by Ferrero Mortara et al. (1984) is Type locality: “… unteren Obermiocän – den Sylvana- similar to the Recent Janulus stephanophorus (Deshayes, 1850) Schichten – von Mörsingen.” and the Caspreno species. Status: uncertain. Probably not a Janulus, but a punctid or a discid species. According to Lueger (1981), the specimen assigned Janulus austriacus Harzhauser & Binder, 2004: 23, Pl. 7, figs 12-15. to this species by Schlickum (1978: Pl. 19, fig. 16) belongs toDiscus Type locality: “Richardhof (sample RH A/2) close to Mödling pleuradras (Bourguignat, 1881). in Lower Austria (Vienna Basin); marls deposited in a Pannonian (late Miocene) swampy lake adjacent to Lake Pannon; mammal Patula (Janulus) olisipponensis Roman, 1907: 82-3, text-fig. 8. zone MN9.” Type locality: “Casaes de Valle d’Obidos près Rio Maior. – Status: probably a Janulus species: although no information Etage Pontique.” is available on radial rows of palatal teeth, it is very similar to the Status: probably a Janulus species: although no information Recent Janulus stephanophorus (Deshayes, 1850). is available on radial rows of palatal teeth, it is very similar to the Recent Janulus stephanophorus (Deshayes, 1850). Janulus decorosus Yü & Zhang, 1982: 64-5, Pl. 7, figs 4-6. Type locality: Baoyue formation, Sanshui Basin, Guangdong; Helix rysa Wood, 1848: 4, Pl. 1, figs 1a-c. Middle Eocene. Type locality: “Red Crag, Walton Naze.” Status: uncertain. Based on very poorly preserved specimens. Status: uncertain. First assigned to Janulus by Sandberger (1875), it recalls J. bifrons from which it differs by virtue of a Patula densestriata Klika, 1891: 40, figs 32a-c. wider umbilicus, more reflexed peristome and blunt peripheral keel Type locality: “bei Wärzen” (Bohemia). (absent in J. bifrons). Status: uncertain. Probably a Janulus species: although no information is available on radial rows of palatal teeth, it is very Janulus schottleri Wenz, 1922: 48, text-figs 1a-d. similar to Janulus schottleri Wenz, 1922. Type locality: “Häufig in den Süsswasserschichten am Streitberg bei Treis a. d. Lumda und bei Ilschhausen.” Helix disparilis Deshayes, 1863: 818, Pl. 52, figs 12-15. Status: very similar to Janulus stephanophorus (Deshayes, Type locality: “Côte Saint-Martin, près d’Etampes … Calcaire 1850). The species has radial row of palatal teeth at 90 degrees to de Beauce.” each other; each row consisting of three teeth: adaxial and medial Status: uncertain. Probably not a Janulus, but a discid species. teeth in the form of a short fold; abaxial larger and V-like. Apparently very similar to Patula densestriata Klika, 1891. Janulus germainae Schlickum & Geissert, 1980: 237-8, Pl. 14, fig. 31. Helix striata Eichwald, 1830: 215, 300 (depicted by Eichwald, Type locality: “Oberes Pliozän, Triptychia-Mergel … 1852: Pl. 11, figs 9a-d). Kiesgrube Mary Kocher, Sessenheim, Unterelsass.” Type locality: “près de Holowczynce, aux environs de Status: uncertain. Probably not a Janulus, but a discid species. Mendzibosh, dans un calcaire tertiaire d’eau douce” (Eichwald, Niederhöfer & Falkner (2004) stated that the type material could 1853). include more than one species. Note: junior primary homonym of Helix striata Müller, 1774, now striata (Pulmonata, ). Patula gottschicki Jooss, 1912: 32-3, Pl. 2, fig. 2. Status: uncertain. Eichwald’s (1852-53) description and figures Type locality: “Pharion’s Sandgrube in Steinheim, Pupaschicht.” may be consistent with a Janulus species, but re-examination is Status: uncertain. Jooss’s description and figure are consistent needed. with a Janulus species (especially due to palatal teeth), but re- examination is needed. G. Manganelli et alii - A new species of Janulus from Zanclean of Tuscany 173

Patula (Janulus) supracostata Sandberger, 1872: Pl. 29, figs 2a-2c (1872), representing the holotype, cannot be identified. However (described by Sandberger, 1875: 584). they thought that it may belong to the “Discus-convergent Janulus Type locality: “Vermes bei Delsberg (Ct. Bern), Birk bei germainae group”. Mörsingen (v. Zell) Georgsgemünd und Hasenmühle bei Eichstädt (Gümbel) im Kalke mit Helix sylvana, Undorf bei Regensburg in Helix suttonensis Wood, 1872: 2, Pl. 1, figs 2a-c. gleichaltem Braunkohlen-Thone (Gümbel), an beiden Orten selten” Type locality: “Coralline Crag, Sutton.” (Sandberger, 1875). Status: probably a Janulus species: although no information Status: uncertain. Sandberger’s description and figure may be is available on radial rows of palatal teeth, it is very similar to the consistent with a Janulus species, but re-examination is needed. Recent Janulus stephanophorus (Deshayes, 1850). According to Moser et al. (2009) the specimen figured by Sandberger

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