The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake (Most Basin) MATHIAS HARZHAUSER, THOMAS A. NEUBAUER, ELISAVET GEORGOPOULOU & JOSEF HARL We present a critical evaluation of the Early Miocene terrestrial and aquatic mollusc fauna of the North Bohemian Lake in the Most Basin in the Czech Republic. In total, 90 species (8 aquatic and 81 terrestrial gastropods, one bivalve) are documented from that lake system that had formed within the North Bohemian Rift. Only three of these species are newly recorded for the Most Basin, suggesting that the fauna is well sampled. Based on historical collections of the Natural History Museum in Vienna, which were partly acquired by quantitative bulk samples from Tuchořice, a rough estimate of the composition of the terrestrial assemblage can be presented. More than 80% of the >30,400 shells are represented by carychiids, vertiginids and valloniids, suggesting the presence of densely forested wetlands fringing the North Bohemian Lake. About 57% of the terrestrial species are known so far exclusively from the Most Basin. This high degree of “endemism”, however, is rather a result of the still very fragmentary knowledge of coeval European faunas. Discus rasseri Harzhauser, Neubauer & Georgopoulou sp. nov. and Discus zagorseki Harzhauser, Neubauer & Georgopoulou sp. nov. are described as new species and Esuinella Harzhauser, Neubauer & Georgopoulou gen. nov. (Valloniidae), Nordsieckula Harl & Harzhauser gen. nov. (Orculidae), and Manganellia Harzhauser, Neubauer & Georgopoulou gen. nov. (Discidae) are introduced as new genera. Serrulastra (Serruplica) tuchoricensis nom. nov. is proposed as replacement name for Clausilia laevigata Frankenberger, 1914. • Key words: Gastropoda, Bivalvia, Miocene, Burdigalian, terrestrial ecosystems, Most Basin. HARZHAUSER, M., NEUBAUER, T.A., GEORGOPOULOU, E. & HARL, J. 2014. The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake (Most Basin). Bulletin of Geosciences 89(4), 819–908 (18 figures, 1 table). Czech Geological Survey, Prague. ISSN 1214-1119. Manuscript received February 10, 2014; accepted in revised form May 5, 2014; published online September 19, 2014; issued September 30, 2014. Mathias Harzhauser (corresponding author), Thomas A. Neubauer, Elisavet Georgopoulou & Josef Harl, Geological-Palaeontological Department, Natural History Museum Vienna, Burgring 7, A-1010 Vienna, Austria; mathias.harzhauser@nhm-wien.ac.at; thomas.neubauer@nhm-wien.ac.at; elisavet.georgopoulou@nhm-wien.ac.at; josef.harl@nhm-wien.ac.at The terrestrial and lacustrine mollusc assemblages from the Most Basin in Bohemia comprise one of the most diverse Early Miocene (Burdigalian) non-marine mollusc faunas of Europe. Many extinct genera are based on species from Tuchořice and numerous authors referred to species from this locality when describing other European Cainozoic mollusc faunas. The exceptionally well-preserved fauna was initially described by Reuss in Reuss & Meyer (1849a, b). During the next decades additions and revisions were presented by Reuss (1861, 1868), Slavík (1869a, b), Boettger (1870a, 1877), Sandberger (1875), and Klika (1890). A first comprehensive synopsis of this pioneer phase was given by Klika (1891, 1892), followed by additional contributions by Babor (1897), Flach (1889, 1891), Frankenberger (1912, 1914), Wenz (1915a), Thuma (1916, 1922) and Petrbok (1925). The wealth of data and references were summarised again by Wenz (1923–1930). Since then no comprehensive evaluation of the fauna was performed. Instead, many DOI 10.3140/bull.geosci.1503 authors discussed certain taxa – often with focus on other regional faunas or addressing specific taxonomic problems – introducing numerous new combinations (e.g. Pfeffer 1930; Wenz & Zilch 1959–1960; Schlickum 1968, 1970a, b; Schlickum & Truc 1972; Nordsieck 1972, 1981a, b, 1986; Schlickum & Strauch 1975; Strauch 1977; Moayedpour 1977; Falkner 1974, 1986; Richardson 1980; Prisyazhnyuk 1984; Čejchan 1985; Boeters et al. 1989; Kadolsky 1993; Stworzewicz 1999a, b; Esu 1999; Manganelli & Giusti 2000a; Kadolsky & Piechocki 2000; Kókay 2006; Schnabel 2006; Binder 2008). This kind of information is often hidden in the publications as the relation to the Most Basin faunas is not always obvious from the titles. Moreover, only a very small percentage of the species has been depicted so far by photographs and SEM-micrographs. We present a synopsis of the Early Miocene molluscs from the North Bohemian Rift. We aim for an updated documentation and illustration of the fauna but do not consider this paper a critical revision of all taxa. 819 Bulletin of Geosciences • Vol. 89, 4, 2014 Figure 1. Position of the North Bohemian Rift in Czech Republic with distribution of Lower Miocene fluvial and lacustrine deposits in the Chomutov and Most basins (after Kvaček et al. 2004) and position of the most important mollusc-bearing localities: Tuchořice (T), Lipno (L), Korozluky (K) and Pyšná (P). For much more detailed informations on regional geology and geographic maps see Heissig & Fejfar (2007). Material The studied material derives mainly from the collections of the Natural History Museum in Vienna (NHMW). This collection comprises a large suite of shells that were bought during the “gold rush” phase of investigations of the fauna from the local Czech teacher Karl Ihl in the late 1880s. Additional collections were bought during this time from Oskar Ritter von Troll-Obergfell, Josef Florian Babor, and Franz Thuma. Only few species, which were listed by Klika (1891) and others from the Most Basin, are missing in the NHMW collections; these species were studied in the collections of the National Museum in Prague (NM-PM-P) and the Senckenberg Naturmuseum in Frankfurt (SMF). All illustrated and studied NHMW material derives from Tuchořice, if not stated otherwise. Geological setting and stratigraphy The Early Miocene mollusc fauna of the Most Basin was collected during the late 19th century at the four villages Tuchořice, Lipno (= Groß Lippen), Korozluky, and Pyšná (= Stolzenhahn) in northern Bohemia (Czech Republic) [the fauna from Dvérce (= Wärzen), as described by Klika (1891), is of Rupelian age (Fejfar 1987, Heissig & Fejfar 2013) and is not considered herein]. The villages Tuchořice and Lipno are situated about 9 km south east of Žatec; Korozluky is about 2 km east of Most; the “Stolzenhahn” 820 locality was situated near Pyšná about 11 km west of Most (see Heissig & Fejfar 2007 for detailed geographic and geologic maps of the area and Klika 1891, Thuma 1922 and Boeters et al. 1989 for data on “Stolzenhahn”). Additional material was detected in drillings in the area of Chomutov (Čtyroký et al. 1964). All localities belong to the 870 km2 large Most Basin, which is part of the North Bohemian Rift (= Ohře-Eger Rift). This tectonic structure formed as result of volcanotectonic subsidence and is delimited by two roughly SW-NE trending tectonic lines (Fig. 1) (Kasiński 1991, Fejfar et al. 2003). Sedimentation in the rift occurred in several sedimentary cycles from Eocene to Plio-Pleistocene times (Suhr 2003). In the Most Basin, the Early Miocene cycle is characterised by thick brown coal seams (“main seam member”) along with fluvial and lacustrine sediments and volcanogenic deposits attaining up to 700 m thickness (Fejfar et al. 2003, Kvaček et al. 2004, Grygar & Mach 2013). Dark grey calcareous marls in the base of the main seam contained a limnic mollusc fauna with Staadtiellopsis rubeschi, which was also detected in drillings close to Chomutov (Čtyroký et al. 1964). Slightly younger aquatic and terrestrial mollusc faunas were collected from bedded limestones, which are exposed on isolated fault blocks along the southern margin of the Most Basin (Tuchořice, Lipno) (Fejfar & Sabol 2005). Closeby, travertines that formed along tectonically induced hot thermal springs are devoid of molluscs (Fejfar et al. 2003). Poisonous CO2-exhalations are discussed as trigger for the Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake considerable taphonomic bias towards carnivorous vertebrates (Mlíkovský 2002, Fejfar et al. 2003, Fejfar & Sabol 2005) and might also explain the absence of terrestrial molluscs. The rich vertebrate fauna of Tuchořice and several other localities in the Most Basin allows a correlation with the European Mammal Zone MN3b, corresponding to the Early Burdigalian (= Eggenburgian) and an absolute age of ~19–18 Ma (Fejfar et al. 2003; Kvaček et al. 2004; Heissig & Fejfar 2007, 2013). Palaeogeography and palaeoecology Kvaček et al. (2004) coined the term “North Bohemian Lake” for the Early Miocene wetlands of the Most Basin and estimated an original extent of roughly 1700 km2 based on the erosional relics of the lake deposits. It was initiated during the Burdigalian by increasing subsidence of the basin, resulting in the formation of swamps and shallow lakes. The North Bohemian Lake was an open system, fed from the south by a stream which entered the basin at Žatec. Towards the north, streams drained into the North Sea Basin. Outflow is considered to have been much lower and evaporation kept the lake level stable (Suhr 2003, Kvaček et al. 2004). The excellent palaeobotanic record of the area suggests dense riparian forests with maples and palms in the Tuchořice area with high precipitation and a mean annual temperature of >18 °C (Fejfar et al. 2003). The lowlands of central and northern Florida were discussed as modern counterpart by Fejfar et al. (2003), whilst Kvaček et al. (2004) proposed the wetlands of the Okavango River in Botswana as modern analogues. A compilation of the manifold fauna and flora of the Miocene Most Basin with numerous environmental reconstructions and a detailed summary on the investigation history is given in Dvořák et al. (2010). Discussion Composition of the fauna In total, 81 terrestrial and 8 aquatic gastropod species and 1 bivalve species are known so far from the Most Basin (Table 1). The NHMW collection – comprising more than 30,400 terrestrial and >30,000 aquatic shells – contains 68 of the terrestrial species, whilst 11 species described by previous authors could not be detected in this “sample” but were available at the collection of the National Museum in Prague. Only three species are documented for the first time from the Most Basin: Punctum propygmaeum Andreae, 1904, Vitrea cf. procrystallina (Andreae, 1902), and Discus rasseri sp. nov. The latter one was lumped with other species and the small Punctum and Vitrea were probably overlooked due to the small size and the lack of SEM-techniques. This suggests that the fauna is well studied and is a reliable example of an Early Miocene terrestrial gastropod assemblage during the Mammal Zone MN3. The huge number of shells and the quantitative sampling method applied by Oskar Ritter von Troll-Obergfell, who washed and sieved large amounts of sediment picking out all shells, allow also to estimate the quantitative composition of the assemblage. Regarding individual numbers, more than 70% of the terrestrial assemblage is represented by only two tiny species: Carychiella eumicrum (Bourguignat, 1857) (~43%) and Vertigo callosa (Reuss in Reuss & Meyer, 1849) (~28%). These are followed by Spermodea plicatella (Reuss in Reuss & Meyer, 1849) (~6%), Esuinella nana (Braun in Walchner, 1851), and Negulus suturalis (Sandberger, 1858) (both ~3%); Acicula fusca frici (Flach, 1889), Klikia labiata (Klika, 1891), and Discus euglyphus (Reuss in Reuss & Meyer, 1849) range between 1–2%. The remaining 71 species each comprise distinctly less than 1% in individual numbers. This pattern is indicative for forested wetlands. Modern carychiids inhabit permanently moist environments such as riparian zones, meadows, and swamps (Weigand & Jochum 2010). Recent Spermodea species are typically found in deciduous woodlands, in humid leaf litter and need continuously moist conditions (Neubert 2011, Welter-Schultes 2012). Forest leaf litter is also the habitat of the African species of Negulus (van Bruggen 1994). The freshwater fauna in turn is low diverse. It is dominated by pulmonates, including one species of each Radix, Stagnicola, Acroloxus, Gyraulus, Hippeutis, and Planorbarius. Concerning abundance, Gyraulus dealbatus (Braun in Walchner, 1851) is by far the most common with several ten thousands of shells, followed by the frequent Planorbarius cornu (Brongniart, 1810). Such a composition is today typically found in stagnant to lentic, highly vegetated environments (Glöer 2002). In contrast, prosobranch gastropods are rare, including Bythinella? scalaris (Slavík, 1869) and the rissooid Staadtiellopsis rubeschi (Reuss in Reuss & Meyer, 1849). The latter species probably derives from a different stratum or area in the lake, as it is not found in our material. There is nothing known about the ecology of the fossil genus Staadtiellopsis. As even its systematic position is still doubtful (variably affiliated with the families Amnicolidae, Micromelaniidae, Pomatiopsidae, and Truncatellidae; Kadolsky 1993, 2008; Kadolsky & Piechocki 2000), it is impossible to infer its ecological preferences. Staadtiellopsis rubeschi appears in huge numbers at the open cast mine Merkur North, where it is always associated with planorbids and lymnaeids (as seen in the illustration in Heissig & Fejfar 2013). Therefore, a preference for lentic conditions may be deduced. 821 Bulletin of Geosciences • Vol. 89, 4, 2014 Taxa indicating riverine influx, like Theodoxus or Ancylus, are absent. Concluding, the composition of the studied assemblages confirms the palaeoenvironmental reconstruction of the North Bohemian Lake as extensive wetlands proposed by Kvaček et al. (2004). Biostratigraphic and biogeographic significance of the fauna Our knowledge of the compositions and interrelations of the European Neogene terrestrial gastropod faunas is still strongly influenced by the milestone catalogus of Wenz (1923–1930). Despite the enormous value of his compilation, it should not be overlooked that many of his references listed in the synonymies have been compiled uncritically. Therefore, historical misidentifications and species-lumping resulted in geologically long-lived and geographically wide-spread “species”, distorting biostratigraphic and biogeographic patterns. Based on the data compiled by Wenz (1923–1930), the Burdigalian fauna from the Most Basin suggested tight relations with the roughly coeval assemblage from Theobaldshof/Rhön and the Chattian assemblages of the Mainz Basin and comparable links to the Middle Miocene, such as those from Opole in Poland and from Rein in Austria. During the last 20 years, new data for several old and new key localities of European terrestrial mollusc faunas have become available (e.g. Moayedpour 1977; Stworzewicz 1993, 1995, 1999a, b; Binder 2002, 2004; Kókay 2006; Salvador 2013; Harzhauser et al. 2014). These data facilitate a critical evaluation of alleged occurrences of Most Basin taxa in other faunas. Based on this survey, the number of shared species of these faunas with those of the Most Basin turned out to be low. Only ~14% of the species occur already in the late Oligocene fauna of Hochheim and only 5–8% of the species occur in the Middle Miocene faunas of Rein and Opole. Even more striking is the rather low percentage of shared species with other Early Miocene faunas. Only 11–15% of the species contribute to the assemblages of Somlóvásárhely (Hungary), Bełchatów (Poland), and Theobaldshof/Rhön (Germany); faunistic relations to the mid- and late Burdigalian faunas of Oberdorf and Teiritzberg in Austria are nearly absent. In total, 45 species (57%) of the terrestrial fauna are known so far exclusively from the Most Basin and only 34 species (43%) are known from other regions as well. For the freshwater fauna, the percentage of endemic taxa is much lower, including only 3 of 9 species (Staadtiellopsis rubeschi is treated as endemic although it is also known from the Cheb and Chomutov basins. These basins, however, were part of the North Bohemian Rift). These observations suggest two conclusions: 1. The turn-over rates in the terrestrial mollusc faunas 822 during the Oligocene and Miocene were probably as high as those of the vertebrate faunas but the lack of critical revisions of nearly all important faunas does not yet allow serious estimates. This assumption is in good agreement with the patterns documented for the Clausiliidae. Due to the detailed investigations by Nordsieck (1972, 1981a, b, 2000), the door snails are currently the only well-studied Neogene group of terrestrial gastropods in Europe and turned out to be stratigraphically highly significant. 2. Even within time slices, the faunas were not homogenous at all. The surprisingly low similarities between the Most Basin assemblages and roughly coeval ones may largely reflect different palaeoecological settings, such as swamps/backswamps (e.g. Oberdorf), carbonate-dominated forested wetlands (e.g. Tuchořice), and coastal marshlands (e.g. Teiritzberg). Conclusions The terrestrial and aquatic mollusc fauna of the Most Basin is among the most important records of the Early Miocene and comprises 90 species. The aquatic assemblages are dominated by planorbids in individual numbers; carychiids, vertiginids, and valloniids characterise the terrestrial assemblages. The composition suggests the presence of densely forested wetlands surrounding a carbonate-dominated lake system, which is in accordance with previous interpretations based on palaeobotanical records (Fejfar et al. 2003). Many species from the Most Basin have been reported also from other Early and Middle Miocene localities of Europe. Most of these turned out to be based on misidentifications and even those accepted herein will need confirmation in many cases. This critical survey reveals that the alleged long stratigraphic ranges of many terrestrial gastropods – which would be in complete opposition to the fast turnover rates observed in Miocene mammal faunas – are largely artefacts due to inadequate taxonomy. Systematic palaeontology The systematic arrangement follows largely Nordsieck (1986, 1987), Hausdorf (2000), Bouchet et al. (2005), and Welter-Schultes (2012). We present a critical chresonymy with main focus on citations referring to occurrences from the Most Basin. The references are reproduced strictly as originally provided by the authors without modification; text in square brackets indicates text that was obviously intended by the author but not fully written at the cited point; e.g. the abbreviation H. when clearly used for Helix is referred to as H.[elix] in the chresonymy. This strict use may be of importance in cases of problematic priority, available combinations, and homonyms. Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake Table 1. List of all Early Miocene mollusc species from the Most Basin accepted herein. Specimen numbers refer to the NHMW collection. No. Species Specimens No. Species Specimens 1 Bythinella? scalaris (Slavík, 1869) 0 47 Serrulastra (Serrulastra) amphiodon (Reuss, 1861) 2 Staadtiellopsis rubeschi (Reuss in Reuss & Meyer, 1849) 0 48 Serrulastra (Serrustigma) polyodon (Reuss, 1861) 5 3 Craspedopoma leptopomoides (Reuss, 1868) 9 49 Serrulastra (Serruplica) tuchoricensis nov. nom. 0 4 Acicula fusca frici (Flach, 1889) 21 571 50 Laminifera mira (Slavík, 1869) 5 5 Acicula limbata Reuss, 1861 11 51 Baboria slaviki (Babor, 1897) 0 6 Platyla callosa (Boettger, 1870) 51 52 Constricta tenuisculpta (Reuss, 1861) 7 Platyla subfusca (Flach, 1889) 8 Radix subovata (von Zieten, 1832) 9 Stagnicola subpalustris (Thomä, 1845) 11 Gyraulus dealbatus (Braun in Walchner, 1851) 53 Cochlodina (Miophaedusa) perforata (Boettger, 1877) 0 9 54 Canalicia attracta (Boettger, 1870) 1 68 10 Acroloxus decussatus (Reuss in Reuss & Meyer, 1849) 1 >10,000 55 Canalicia klikai (Babor, 1897) 0 56 Pseudocalaxis? insignis (Babor, 1897) 0 57 Palaeoglandina gracilis (von Zieten, 1832) 13 Planorbarius cornu (Brongniart, 1810) 432 Pseudoleacina oligostropha (Reuss in Reuss & Meyer, 58 1849) 14 Carychiopsis schwageri (Reuss, 1868) 161 59 12 Hippeutis ungeri (Reuss in Reuss & Meyer, 1849) 1 15 Carychiopsis prisyazhnyuki Stworzewicz, 1999 16 Carychiella eumicrum (Bourguignat, 1857) 3 13,2 17 Azeca pumila Slavík, 1869 8 18 Azeca monocraspedon Slavík, 1869 3 19 Hypnophila subrimata (Reuss in Reuss & Meyer, 1849) 60 20 Cochlicopa dormitzeri (Reuss in Reuss & Meyer, 1849) 5 21 Mastus complanatus (Reuss in Reuss & Meyer, 1849) 63 22 Mastus filocinctus (Reuss, 1861) 2 23 Paracoryna diezi (Flach, 1891) 8 Pseudoleacina producta (Reuss in Reuss & Meyer, 1849) 162 60 Punctum propygmaeum Andreae, 1904 27 62 Discus euglyphus (Reuss in Reuss & Meyer, 1849) 432 63 Discus rasseri sp. nov. 283 64 Discus stenospira (Reuss in Reuss & Meyer, 1849) 65 Discus zagorseki sp. nov. 66 Manganellia alata (Klika, 1891) 0 2 59 70 Aegopinella denudata (Reuss in Reuss & Meyer, 1849) 71 Aegopinella vetusta (Klika, 1891) 27 Pleurodiscus falciferus (Boettger, 1870) 28 Vallonia lepida (Reuss in Reuss & Meyer, 1849) 29 Acanthinula tuchoricensis (Klika, 1891) 31 Esuinella nana (Braun in Walchner, 1851) 32 Spermodea plicatella (Reuss in Reuss & Meyer, 1849) 248 0 17 19 73 Miozonites algiroides (Reuss in Reuss & Meyer, 1849) 60 18 74 Limacus crassitesta (Reuss, 1868) 1 Phenacolimax intermedius (Reuss in Reuss & Meyer, 75 1849) 9 0 843 1,915 33 Granaria intrusa (Slavík, 1869) 22 72 Lyrodiscus ihli (Klika, 1891) 202 30 Acanthinula sp. 3 17 67 Janulus sp. 25 Strobilops (Eostrobilops) elasmodonta (Reuss, 1861) 201 6 68 Vitrea cf. procrystallina (Andreae, 1902b) 69 Oxychilus mendicus (Slavík, 1869) Strobilops (Discostrobilops) uniplicatus (Braun in Walchner, 1851) 102 61 Discus bohemicus (Wenz in Fischer & Wenz, 1914) 44 26 11 151 Strobilops (Eostrobilops) fischeri (Wenz in Fischer & Wenz, 1914) 24 11 4 13 76 Phenacolimax crassitesta (Andreae, 1902) 0 77 Leucochroopsis apicalis (Reuss, 1861) 140 78 Pseudomonacha zippei (Reuss in Reuss & Meyer, 1849) 150 1 79 Pseudomonacha homalospira (Reuss, 1861) 83 35 Vertigo angulifera Boettger, 1884 0 31 36 Vertigo minor Boettger, 1870 3 Praeoestophorella petersi (Reuss in Reuss & Meyer, 80 1849) 81 Protodrepanostoma involuta nordsiecki Falkner, 1986 87 34 Ptychalaea flexidens (Reuss, 1861) 37 Vertigo callosa (Reuss in Reuss & Meyer, 1849) 38 Vertigo tuchoricensis Pilsbry in Pilsbry & Cooke, 1919 39 Negulus suturalis (Sandberger, 1858) 40 Negulus raricosta (Slavík, 1869) 41 Gastrocopta (Albinula) turgida (Reuss in Reuss & Meyer, 1849) 42 Truncatellina splendidula (Sandberger, 1875) 43 Nordsieckula subconica (Sandberger, 1858) 44 Oxyloma affinis (Reuss in Reuss & Meyer, 1849) 45 Triptychia vulgata (Reuss in Reuss & Meyer, 1849) 46 Serrullella schwageri (Boettger, 1877) 8,6 1 842 7 106 268 1 128 13 2 82 Protodrepanostoma hecklei (Klika, 1891) 6 83 Metacampylaea papillifera (Klika, 1891) 0 Pseudochloritis robusta (Reuss in Reuss & Meyer, 84 1849) 52 85 Klikia labiata (Klika, 1891) 447 86 Apula devexa (Reuss, 1861) 60 87 Apula prominens (Babor, 1897) 88 Creneatachea obtusecarinata (Sandberger, 1858) 89 Megalotachea macrocheila (Reuss in Reuss & Meyer, 1849) 90 Sphaerium prominulum (Reuss in Reuss & Meyer, 1849) 0 151 175 15 823 Bulletin of Geosciences • Vol. 89, 4, 2014 Most of the taxa have been described in great detail by Klika (1891). This milestone paper is freely available online. Therefore, we will not repeat all the descriptions but focus mainly on taxonomic problems and distribution data. hydrobiid, which cannot be identified at all. Bythinella species are bound to springs and rarely to caves and groundwater (Bichain et al. 2007). Therefore, its presence at the springs of Tuchořice is not unrealistic. Unfortunately, the descriptions of Slavík (1869a, b) do not allow any generic identification. Class Gastropoda Cuvier, 1795 Subclass Orthogastropoda Ponder & Lindberg, 1995 Superorder Caenogastropoda Cox, 1960 Order Neotaenioglossa Haller, 1892 Occurrence. – Only known from Tuchořice. Superfamily Rissooidea Gray, 1847 Family Amnicolidae Tryon, 1863 Genus Bythinella Moquin-Tandon, 1856 Type species. – Bulimus viridis Poiret, 1801; subsequent designation by ICZN Opinion 2161 (Case 3321). Recent, France. Bythinella? scalaris (Slavík, 1869) species inquirenda *1869a Paludinella scalaris sp. nov.; Slavík, p. 267, pl. 4, figs 24, 25. 1869b Paludinella scalaris sp. nov.; Slavík, p. 269, pl. 4, figs 24, 25. 1891 Bythinella scalaris Slavík. – Klika, p. 115, text-figs 114a, b. 1892 Bythinella scalaris Slavík. – Klika, p. 110, text-figs 114a, b. 1911 Bythinella scalaris Sl. – Kafka, p. 69. 1917 Bythinella scalaris Slavik. – Wenz, p. 77. 1926 Bythinella (Bythinella) scalaris (Slavik). – Wenz, p. 2030 (cum syn.). Material. – No material available. Dimensions (after Klika 1891). – Diameter: 0.75 mm, height: 2.5 mm. Discussion. – We have tried to find the type specimen in the collection of the National Museum in Prague. Klika (1891) mentions a single well preserved shell. The available specimen, however, is a poorly preserved shell of a Superfamily Truncatelloidea Gray, 1840 Family Emmericiidae Brusina, 1870 Genus Staadtiellopsis Schlickum, 1968 Type species. – Cyclostoma rubeschi Reuss, 1849; original designation. Early Miocene, Czech Republic. Staadtiellopsis rubeschi (Reuss in Reuss & Meyer, 1849) Figure 2A, B, E, F, M 1849a Cyclostoma Rubeschi Rss.; Reuss in Reuss & Meyer, p. 12 (nomen nudum). *1849b C.[yclostoma] Rubeschi m.; Reuss in Reuss & Meyer, p. 40, pl. 4, fig. 12. 1861 C.[yclostoma] (Pomatias) Rubeschi Reuss. – Reuss, p. 62. 1875 Euchilus? Rubeschi Reuss. – Sandberger, p. 423, pl. 24, figs 31–31b. 1891 Euchilus Rubeschi Reuss. – Klika, p. 113, text-figs 112a, b. 1892 Euchilus Rubeschi Reuss. – Klika, p. 108, text-figs 112a, b. 1911 Euchilus rubeschi Rss. – Kafka, p. 69. 1916 Euchilus Rubeschi Reuss. – Thuma, p. 84. 1917 Staliola rubeschi (Reuss). – Wenz, p. 77. 1923 Staliola (Staliola) rubeschi (Reuss). – Wenz, p. 2189. 1964 Nystia rubeschi (Reuss, 1849). – Čtyroký et al., p. 138, figs 6, 7. 1968 Nystia (Staadtiellopsis) rubeschi (Reuss). – Schlickum, p. 45, pl. 5, figs 10, 11. 1970b Nystia (Staadtiellopsis) rubeschi (Reuss). – Schlickum, p. 292, fig. 14. 1993 Staadtiellopsis rubeschi (Reuss, 1849) sensu Schlickum, 1968. – Kadolsky, p. 385, fig. 92. Figure 2. A – Staadtiellopsis rubeschi (Reuss in Reuss & Meyer, 1849), NM-PM-P 1964. • B – Staadtiellopsis rubeschi (Reuss in Reuss & Meyer, 1849), NM-PM-P 1965. • C – Craspedopoma leptopomoides (Reuss, 1868), NHMW 2013/0572/0001. • D – Craspedopoma leptopomoides (Reuss, 1868), NHMW 2013/0572/0001.• E, F – Staadtiellopsis rubeschi (Reuss in Reuss & Meyer, 1849), NM-PM-P 1966. • G, H – Acicula fusca frici (Flach, 1889), NHMW 2013/0572/0002. • I – Acicula fusca frici (Flach, 1889), NHMW 2013/0572/0002 (sinistral specimen). • J – Acicula limbata Reuss, 1861, NHMW 2013/0572/0003. • K – Acicula limbata Reuss, 1861, NHMW 2013/0572/0003. • L – protoconch of J. • M – Staadtiellopsis rubeschi (Reuss in Reuss & Meyer, 1849), NM-PM-P 1967. • N–P – Platyla callosa (Boettger, 1870), NHMW 2013/0572/0004. • Q, R – Platyla subfusca (Flach, 1889), NHMW 2013/0572/0005. • S – Platyla subfusca (Flach, 1889), NHMW 2013/0572/0005. • T – protoconch of Q, R. 824 Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake C B A E M G N D H O J I Q K S R F L P T 825 Bulletin of Geosciences • Vol. 89, 4, 2014 2000 Staadtiellopsis rubeschi (Reuss, 1849). – Kadolsky & Piechocki, p. 225, figs 15, 16. 2013 Nystia. – Heissig & Fejfar, p. 8, fig. 2b. non 1997 Staadtiellopsis rubeschi (Reuss, 1849) (?). – Piechocki, p. 109, text-figs 6, 7 (= Staadtiellopsis schlickumi Kadolsky & Piechocki, 2000). non? 2006 Staadtiellopsis rubeschi (Reuss), 1849 var. – Kókay, p. 43, pl. 13, figs 13, 14, text-fig. 1. Material. – 5 specimens in the Prague collection (NM-PM-P 1964–1968) from the Merkur-North mine in the Chomutov Basin; no material available in the NHM collection. Dimensions. – Diameter: 2.8 mm, height: 5.5 mm (Fig. 2A); diameter: 3.1 mm, height: 5.2 mm (Fig. 2B). Discussion. – Kadolsky & Piechocki (2000, p. 225) and Kadolsky (2008) gave an overview about the status of this species, which they understand as Staadtiellopsis rubeschi (Reuss, 1849) sensu Sandberger (1875) and Schlickum (1968). An important feature is the complex internal structure of the aperture with a deep groove terminated by a free-ending internal shell layer. The illustration in Klika (1891) is a poor drawing and does not show the characteristic aperture (Čtyroký et al. 1964). The protoconch is bulbous and entirely smooth and consists of ca 1 whorl. Some specimens have a decollate apex. Occurrence. – Only mentioned from Korozluky and Pyšná in the Most Basin. Alleged occurrences at Tuchořice are doubtful. This species is most abundant in the Chomutov Basin. It was detected in all drillings described by Čtyroký et al. (1964) and in marls intercalated in lignite seams of Merkur-North and Ahníkov in the Chomutov Basin, where it is among the dominant species in the lacustrine assemblages. These lignite seams are older than the Tuchořice assemblages (MN3a according to Heissig & Fejfar 2013). Also recorded from Horní Ves (Oberndorf) in the Cheb Basin (Wenz 1923); a further occurrence might be represented in the Burdigalian deposits of the Szcerców lignite in Poland, mentioned without description by Wagner & Matl (2007). Kókay (2006) described this species as “var.” from the late Badenian (Serravallian) of Hungary. These specimens lack the aperture and are much smaller than the Bohemian specimens. In respect to the conchological differences and the large stratigraphic gap of ca 5 Ma between both occurrences, the Hungarian shells are most probably not conspecific with S. rubeschi. Clade Architaenioglossa Haller, 1890 Superfamily Cyclophoroidea Gray, 1847 Family Craspedopomatidae Kobelt & Möllendorff, 1898 826 Genus Craspedopoma Pfeiffer, 1847 Type species. – Cyclostoma lucidum Lowe, 1831; by monotypy. Recent, Madeira. Craspedopoma leptopomoides (Reuss, 1868) Figure 2C, D *1868 Valvata leptopomoides sp. nov.; Reuss, p. 83, pl. 1, fig. 4. 1869a Valvata leptopomoides Reuss. – Slavík, p. 267. 1869b Valvata leptopomoides Rss. – Slavík, p. 270, pl. 4, figs 26–28. 1870a Valvata leptopomoides Rss. – Boettger, p. 299. 1875 Craspedopoma leptopomoides Reuss. – Sandberger, p. 445, pl. 24, figs 33a, b. 1891 Craspedopoma leptopomoides Reuss. – Klika, p. 15, text-fig. 5. 1892 Craspedopoma leptopomoides Reuss. – Klika, p. 15, text-fig. 5. ? 1902a Craspedopoma leptopomoides (Reuss). – Andreae, p. 7. ? 1902b Craspedopoma leptopomoides (Rss.). – Andreae, p. 22. 1911 Craspedopoma leptopomoides Rss. – Kafka, p. 66. 1917 Bolania (Bolania) leptopomoides (Reuss). – Wenz, p. 77. 1923 Bolania (Bolania) leptopomoides (Reuss). – Wenz, p. 1768. Material. – 2 specimens (NHMW 2013/0572/0001), 7 specimens (NHMW 1909/0001/0001). Dimensions. – Diameter: 5.9 mm, height: 5.8 mm (Fig. 2C); diameter: 5.1 mm, height: 5.2 mm (Fig. 2D). Discussion. – This species was traditionally placed in Craspedopoma Pfeiffer, 1847, which is restricted today to the Eastern Atlantic region (Madeira, Azores, Portugal). The absence of any opercula in the fossil record would fit to the corneous nature of the opercula of the Craspedopomatidae. Nevertheless, a detailed revision is needed to decide whether all the Miocene-Pliocene European species, which have been assigned to Craspedopoma, really belog to this genus. Among the Recent species, Craspedopoma hespericum (Morelet & Drouët, 1857) from the Azores is most similar to C. leptopomoides concerning the shell outline, thin peristome, and wide umbilicus. Craspedopoma handmanni Troll, 1907 from the Late Miocene of the Vienna Basin differs in its higher shell and the ovoid aperture. As discussed by Lueger (1981) it might rather represent a Bithyniidae. Wenz (1917, 1923) placed this species in Bolania Gray, 1840, which he considered a senior synonym of Craspedopoma Pfeiffer, 1847. This assignment is incorrect Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake because Bolania Gray, 1840 is a nomen nudum (see Bouchet et al. 2005). The Bohemian shells are reminiscent of the Early Miocene “Procyclotella?” glazeki Stworzewicz, 1995 from Bełchatów in Poland (see note on Procyclotella below). They differ especially in the higher spire and less turbinate outline and lack the typical microscopic spiral sculpture. In addition, the Bohemian species lacks the thickened inner labrum close to the aperture as is characteristic for Procyclotopsis Wenz, 1924. Note on Procyclotella. – Wenz (1924, p. 223) introduced Procyclotopsis as replacement name for Cyclotellina Wenz, 1923a, which is preoccupied by Cyclotellina Cossmann, 1886. Unfortunately, in his corrigenda chapter to his catalogue, Wenz (1930, p. 3040) erroneously wrote Procyclotella, which was used by subsequent authors such as Wenz (1938) and Stworzewicz (1995). Occurrence. – Only known from Tuchořice. Andreae (1902a, b) mentioned this species from Opole (Poland) without description or illustration. These specimens could also represent one of the Craspedopomatidae species described by Stworzewicz (1995) from Bełchatów in Poland. Similarly, Winkler-Hermaden (1951) listed this species from the Ottnangian (~mid-Burdigalian) of Oberdorf (Styrian Basin, Austria), from where it could not be detected again by Binder (2004). Family Aciculidae Gray, 1850 Dimensions. – Diameter: 0.8 mm, height: 1.95 mm (Fig. 2G); diameter: 0.75 mm, height: 2.05 mm (Fig. 2H). Discussion. – Two aciculid species with axial grooves are recorded from the Most Basin: Acicula fusca frici (Flach, 1889) and Acicula limbata Reuss, 1861. As already defined by Flach (1889), both species can be clearly distinguished by the characteristic sutural thread and larger size of A. limbata. Moreover, A. limbata displays a distinctly less convex transition from periphery into base, its aperture is adapically acuminate-elongate, and its protoconch is much lower. Boeters et al. (1989) treat Acme frici Flach, 1889 as a synonym of the Recent Acicula fusca but state that it is more slender. This would make this species a surprisingly long-lived species, which persisted for more than 18 Ma. In our opinion this “genetic stasis” is not very likely and therefore, we propose to keep the slender Bohemian Early Miocene specimens separate at least as chrono-subspecies. Within the 572 specimens, a single sinistral specimen was detected as well (Fig. 2I). Occurrence. – Known from Tuchořice and the Chomutov drillings and from the Aquitanian of Theobaldshof/Rhön in Germany (Moayedpour 1977). A further occurrence from the Lower Oligocene of Detaň (Czech Republic), mentioned by Mikuláš et al. (2003), needs verification but seems unlikely in respect to the very large age difference. Genus Acicula Hartmann, 1821 Type species. – Auricula lineata Draparnaud, 1805; by monotypy. Recent, France. Acicula fusca frici (Flach, 1889) Figure 2G–I *1889 1891 1892 1911 1917 1923 1964 Acme frici n.; Flach, p. 72, fig. 6. Acme friči Flach. – Klika, p. 19, text-fig. 10. Acme friči Flach. – Klika, p. 19, text-fig. 10. Acme Friči Flach. – Kafka, p. 67. Acme (Pupula) frici (Flach). – Wenz, p. 77. Pupula friči (Flach). – Wenz, p. 1856 (cum syn.). Acme friči (Flach, 1889). – Čtyroký et al., table for p. 149. 1977 Acicula (Acicula) frici (Flach). – Moayedpour, p. 57, pl. 1, figs 13, 14. 1989 Acicula fusca (Montagu, 1803). – Boeters et al., p. 43 (pars), text-fig. 41. Material. – 2 specimens (NHMW 2013/0572/0002), 570 specimens (NHMW 1909/0001/0005). Acicula limbata Reuss, 1861 Figure 2J–L 1849a Acme fusca Walk. – Reuss in Reuss & Meyer, p. 11 (non [Turbo] fuscus Montagu, 1803). 1849b A.[cme] fusca Walk. – Reuss in Reuss & Meyer, p. 40, pl. 3, fig. 16 (non [Turbo] fuscus Montagu, 1803). *1861 A.[cicula] limbata Reuss; Reuss, p. 61. 1869a Acicula limbata Reuss. – Slavík, p. 266. 1870a Acicula limbata Rss. – Boettger, p. 284. 1875 Acicula limbata Reuss. – Sandberger, pp. 410, 445, pl. 24, figs 30a, b. 1889 [Acme] limbata Reuss. – Flach, p. 71, fig. 3. 1891 Acme laevissima n.; Klika, p. 18, text-fig. 8. 1891 Acme limbata Reuss. – Klika, p. 18, text-fig. 9. 1892 Acme laevissima Klika. – Klika, p. 18, text-fig. 8. 1892 Acme limbata Reuss. – Klika, p. 18, text-fig. 9. ? 1902b Acme limbata Rss. – Andreae, p. 24. 1911 [Acme] (Tudora) limbata Rss. – Kafka, p. 67. 1911 [Acme] laevissima Kl. – Kafka, p. 67. 1916 [Acme] limbata Reuss. – Thuma, p. 82. 827 Bulletin of Geosciences • Vol. 89, 4, 2014 1917 1922 1923 1923 Acme (Pupula) limbata (Reuss). – Wenz, p. 76. Acme laevissima Kl. – Thuma, p. 5. Acme (Acme) laevissima Klika. – Wenz, p. 1853. Pupula limbata (Reuss). – Wenz, p. 1857 (cum syn.). 1989 Acicula limbata Reuss, 1860. – Boeters et al., p. 56, text-figs 60, 61, non fig. 56. 1989 [Platyla] laevissima. – Boeters et al., p. 217. Material. – One specimen (NHMW 2013/0572/0003), 10 specimens (NHMW 1909/0001/0004). Dimensions. – Diameter: 1.1 mm, height: 2.8 mm (Fig. 2J); diameter: 1.15 mm, height: 2.9 mm (Fig. 2K). Discussion. – Acicula limbata is easily recognised by its deep axial grooves and the typical sutural band along the upper suture of the teleoconch whorls. This species was originally described by Reuss (1861) from the Early Miocene of Bohemia. Nevertheless, Boeters et al. (1989) consider also a Recent species from the Caucasus in Georgia to be conspecific with A. limbata. The only Recent specimen illustrated by Boeters et al. (1989, fig. 56) is stouter, bullet-shaped, the sutural band is broader, its columellar-lip is more concave and the axial grooves are more oblique compared to A. limbata. Therefore, – and also with respect to the huge stratigraphic gap – we doubt that both taxa are conspecific. Klika (1891) described a poorly preserved specimen from Pyšná as Acme laevissima. Based on this description and the very poor illustration, Boeters et al. (1989) placed Acme laevissima in Platyla. Unfortunately, the original description and illustrations are misleading. The type specimen in the collection of the National Museum Prague (NM-PM-P 449) shows axial grooves and a poorly preserved sutural band, which have not been mentioned by Klika (1891). Thus, sculpture, size and overall shape range well within Acicula limbata Reuss, 1861. Acicula crassistoma Stworzewicz & Sołtys, 1996 from the Early/Middle Miocene of Bełchatów was partly lumped with A. limbata but differs clearly in its smaller size and stout shell (Stworzewicz & Sołtys 1996). Occurrence. – Known from Korozluky, Pyšná and Tuchořice. The record from Opole in Poland, mentioned by Andreae (1902b), needs confirmation and might also represent species such as Acicula crassistoma Stworzewicz & Sołtys, 1996. Genus Platyla Moquin-Tandon, 1856 Type species. – Acme dupuyi Paladilhe, 1868; subsequent designation by Boeters et al. (1989). Recent, France. 828 Platyla callosa (Boettger, 1870) Figure 2N–P *1870a Acicula callosa Boettg.; Boettger, p. 284, pl. 13, figs 1a, b. 1889 [Acme] callosa Böttg. – Flach, p. 71, fig. 2. 1891 Acme callosa Boettger. – Klika, p. 16, text-fig. 6. 1892 Acme callosa Boettger. – Klika, p. 16, text-fig. 6. 1911 Acme callosa Bttg. – Kafka, p. 67. 1916 Acme callosa Boettger. – Thuma, p. 82. 1917 Acme (Platyla) callosa Boettger. – Wenz, p. 76. 1923 Acme (Acme) callosa Boettger. – Wenz, p. 1851. 1976 Acicula (Platyla) callosa (Boettger). – Zilch, p. 123, pl. 11, fig. 2. 1989 Platyla callosa (O. Boettger, 1870). – Boeters et al., p. 100, text-figs 92, 94. ? 2006 Platyla callosa (Boettger), 1870 var. – Kókay, p. 44, pl. 14, figs 3, 4 (? = Platyla falkneri Boeters et al., 1986). Material. – 3 specimens (NHMW 2013/0572/0004), 48 specimens (NHMW 1909/0001/0002). Dimensions. – Diameter: 1.4 mm, height: 3.4 mm (Fig. 2N); diameter: 1.35 mm, height: 3.4 mm (Fig. 2O). Discussion. – Platyla callosa (Boettger, 1870) differs from Platyla subfusca (Flach, 1889) in its larger size, it lacks the delicate sutural thread, has an acuminate adapical aperture, develops only a weak cervical callus, is more subcylindrical, and its whorls are less convex. The illustration in Boettger (1870a) is not precise and suggests a bullet-shaped, cylindrical shell, whilst the lectotype in Zilch (1976) has incised sutures and convex whorls. Kókay (2006) reports this species also from the Early Miocene of Hungary based on two fragments of the last whorl. The prominent umbilical callus (sensu Boeters et al. 1989) and the broad external peristomal varix of these specimens might point rather to Platyla falkneri Boeters et al. (1989) as discussed by Kókay (2006). Occurrence. – Only known from Korozluky and Tuchořice. Platyla subfusca (Flach, 1889) Figure 2Q–T *1889 1891 1892 1904 [Acme] subfusca n.; Flach, p. 71, fig. 1. Acme subfusca Flach. – Klika, p. 17, text-fig. 7. Acme subfusca Flach. – Klika, p. 17, text-fig. 7. Acme callosiuscula sp. nov.; Andreae, p. 14, fig. 13. 1911 [Acme] subfusca Flach. – Kafka, p. 67. Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake 1917 Acme (Platyla) subfusca Flach. – Wenz, p. 76. 1923 Acme (Acme) subfusca Flach. – Wenz, p. 1853 (cum syn.). 1976 Acicula (Platyla) callosiuscula Andreae. – Schlickum, p. 3, pl. 1, fig. 4. 1989 Platyla subfusca (Flach, 1889). – Boeters, p. 169, text-figs 170, 171. 2006 Platyla subfusca (Flach), 1889. – Kókay, p. 44, pl. 14, fig. 5. Material. – 2 specimens (NHMW 2013/0572/0005), 2 specimens (NHMW 1909/0001/0003). Dimensions. – Diameter: 1.15 mm, height: 2.9 mm (Fig. 2Q); diameter: 1.2 mm, height: 2.9 mm (Fig. 2S). Discussion. – The most characteristic feature of this species is the prominent external peristomal varix, which is absent in all other aciculids of the Most Basin. Occurrence. – The species was described from the Early Miocene of Tuchořice and from Somlóvásárhely in Hungary (Kókay 2006). It is also reported from the Late Oligocene of Hochheim-Flörsheim in Germany (Wenz 1923). Tentatively, Boeters et al. (1989) considered Platyla callosiuscula (Andreae, 1904) as synonym, which would increase the range of the species to the Middle Miocene (e.g. Zwiefaltendorf, Germany and Opole, Poland). Clade Panpulmonata Jörger et al., 2010 Superorder Basommatophora Keferstein in Bronn, 1864 Order Hygrophila Férussac, 1822 Suborder Branchiopulmonata Morton, 1955 Infraorder Lymnaeoinei Minichev & Starobogatov, 1975 Superfamily Lymnaeoidea Rafinesque, 1815 Family Lymnaeidae Rafinesque, 1815 Genus Radix Montfort, 1810 Type species. – Helix Auricularia Linnaeus, 1758; original designation. Recent, Europe. Radix subovata (von Zieten, 1832) Figure 3A–C, F *1832 Limnaea subovata Hartmann; von Zieten, p. 39, pl. 30, figs 2a, b. 1845 Limnaeus pachygaster Nob.; Thomä, p. 155, pl. 4, fig. 1. 1849a L.[imnaeus] vulgaris Pfr. – Reuss in Reuss & Meyer, p. 1 (non Limnaeus vulgaris Pfeiffer, 1821). 1849b L.[imnaeus] vulgaris Pfr. – Reuss in Reuss & Meyer, p. 37, pl. 4, fig. 6 (non Limnaeus vulgaris Pfeiffer, 1821). 1849a Limnaeus Thomae Rss. – Reuss in Reuss & Meyer, p. 11 (nomen nudum). 1849b L.[imnaeus] Thomae m.; Reuss in Reuss & Meyer, p. 36, pl. 4, fig. 4. 1858 Limnaeus pachygaster Thomæ. – Sandberger, p. 67, pl. 7, figs 1, 4. 1861 L.[imnaeus] pachygaster Thom. – Reuss, p. 78. 1861 L.[imnaeus] Thomaei [sic] Reuss. – Reuss, p. 79. 1875 Limnaeus pachygaster Thomae. – Sandberger, p. 494, pl. 25, fig. 13. 1891 Limnaeus pachygaster Thomae. – Klika, p. 103, text-figs 99a, b. 1891 Limnaeus Thomaei Reuss. – Klika, p. 104, text-fig. 101. 1891 L.[imnaeus] Klikae Bttgr. – Boettger, p. 231. 1892 Limnaeus pachygaster Thomae. – Klika, p. 98, text-figs 99a, b. 1892 Limnaeus Thomaei Reuss. – Klika, p. 100, text-fig. 101. 1911 Limnaeus pachygaster Tho. – Kafka, p. 69. 1915 Limnaea (Limnus) pachygaster Tho. – Fischer & Wenz, p. 56, pl. 2, figs 10a, b. 1916 Limnaeus pachygaster Thomae. – Thuma, p. 84. 1916 [Limnaeus] Thomaei Reuss. – Thuma, p. 84. 1917 Limnaea (Limnaea) pachygaster (Thomae). – Wenz, p. 74. 1923 Radix (Radix) subovata (Zieten). – Wenz, p. 1291. 1923 Radix (Radix) subovata thomaei (Reuss). – Wenz, p. 1310 (cum syn.). 1977 Radix (Radix) subovata (Zieten). – Moayedpour, p. 59. Material. – 2 specimens (NHMW 2013/0572/0006), 3 specimens (NHMW 2013/0572/0007), 2 specimens (NHMW 1909/0001/0085), 2 specimens (NHMW 1909/0001/0087), 1 specimen (NM-PM-P 885). Dimensions. – Diameter: 11.5 mm, height: 19.5 mm (Fig. 3A, F); diameter: 15.5 mm, height: 28 mm (Fig. 3B, C). Discussion. – This species is difficult to distinguish from the co-occurring Stagnicola subpalustris (Thomä, 1845). Both species exhibit a considerable degree of (overlapping) morphological variability, making a strict delimitation almost impossible. Generally, R. subovata has a broader and higher last whorl, producing a more bulbous appearance, with a strong columellar fold, while S. subpalustris has a slender shape and one additional whorl. Each of these characters varies strongly among the studied material and might also depend on the individual age. As both forms have been cited from the same localities in the Most Basin (except Lipno, 829 Bulletin of Geosciences • Vol. 89, 4, 2014 where only S. subpalustris was recorded), they might turn out to reflect one highly polymorphic species (as is known for lymnaeids in general; e.g. Glöer 2002). The specimens, which were identified as “Limnaeus Thomae” by Reuss in Reuss & Meyer (1849) and ranked as subspecies of R. subovata by Wenz (1923), range fully within the variability of R. subovata. This regards especially the slightly inflated last whorl. As any delimitation appears arbitrary we synonymise these records. Occurrence. – Recorded for Korozluky, Tuchořice, and Pyšná in the Most Basin. Additionally recorded for the Lower Miocene of the Aquitaine Basin, the Mainz and Hanau basins, the Rhön Mountains, Southern Germany, and Switzerland (Wenz 1923, Moayedpour 1977). Genus Stagnicola Jeffreys, 1830 Type species. – Lymnaea communis Jeffreys, 1830, by monotypy. Recent, Europe. Stagnicola subpalustris (Thomä, 1845) Figure 3D, E *1845 Limnæus subpalustris Nob.; Thomä, p. 156, pl. 4, figs 9a, b. 1845 Limnæus minor Nob.; Thomä, p. 157. 1849a Limaeus acutus Braun. – Reuss in Reuss & Meyer, p. 11. 1849a L.[imnaeus] medius Rss. – Reuss in Reuss & Meyer, p. 11 (nomen nudum). 1849b L.[imnaeus] acutus Braun. – Reuss in Reuss & Meyer, p. 35, pl. 4, fig. 3. 1849b L.[imnaeus] medius m.; Reuss in Reuss & Meyer, p. 36, pl. 4, fig. 5. 1858 Limnaeus subpalustris Thomæ. – Sandberger, p. 68, pl. 7, figs 2–2b. 1858 Limnaeus minor Thomæ. – Sandberger, p. 70, pl. 7, figs 6–6b. 1861 L.[imnaeus] subpalustris Thom. – Reuss, p. 78. 1861 L.[imnaeus] minor Thom. – Reuss, p. 79. 1875 Limnaeus subpalustris Thomae. – Sandberger, p. 495, pl. 25, fig. 14. 1878 L.[imneus] subpalustris Tho. – Boettger, p. 2. 1878 Limneus minor Tho. – Boettger, p. 4. ? 1878 Limneus Dupuyanus Noulet. – Boettger, p. 5 (non Limnaea Dupuyana Noulet, 1854). 1891 Limnaeus subpalustris Thomae. – Klika, p. 104, text-figs 100a, b. 1891 Limnaeus minor Thomae. – Klika, p. 99, text-figs 102a, b. 1892 Limnaeus subpalustris Thomae. – Klika, p. 104, text-figs 100a, b. 830 1892 Limnaeus minor Thomae. – Klika, p. 101, text-figs 102a, b. 1911 [Limnaeus] minor Th. – Kafka, p. 69. 1916 [Limnaeus] subpalustris Thomae. – Thuma, p. 84. 1916 [Limnaeus] minor Thomae. – Thuma, p. 84. 1917 Limanea (Limnophysa) subpalustris (Thomae). – Wenz, p. 50, p. 74. 1923 Galba (Galba) subpalustris subpalustris (Thomae). – Wenz, p. 1398. 1923 Galba (Galba) subpalustris minor (Thomae). – Wenz, p. 1403 (cum syn.). 1977 Stagnicola (Stagnicola) subpalustris (Thomae). – Moayedpour, p. 59, pl. 2, fig. 6. Material. – One specimen (illustrated, NHMW 2013/0572/0008), 62 specimens (NHMW 2013/0572/0009), 2 specimens (NHMW 1909/0001/0086), 3 specimens (NHMW 1909/0001/0088), 3 specimens (NHMW 1890/0013/0416). Dimensions. – Diameter: 11.3 mm, height: 25.3 mm. Discussion. – Reuss (1861) mentioned L. minor Thomä, 1845 as rare element in Tuchořice. This record as well as the status of this species, which was described from the Early Miocene of the Mainz Basin, is doubtful. It was not illustrated and only poorly described by Thomä (1845), who did not discuss any differences from other cooccurring lymnaeids. Sandberger (1858) and Boettger (1878) depicted small ovoid shells that roughly correspond to Thomä’s description. Yet these specimens are apparently juvenile shells and do not differ from comparable ontogenetic stages of herein studied specimens of S. subpalustris. The “always distinctly lipped peristome” characterising L. minor after Boettger (1878), is also found in several of our subadult specimens. Therefore, we synonymise both taxa. Boettger (1878) determined some specimens from Tuchořice as L. Dupuyanus Noulet. Such ovoid shells with small aperture and thickened peristome are not found in our material. As this species was originally described by Noulet (1854) from the Middle Miocene of Sansan (France), this record is probably a misidentification and is tentatively synonymised with S. subpalustris. For differences to the co-occurring Radix subovata see discussion above. Occurrence. – Known from Korozluky, Tuchořice, Pyšná, and Lipno in the Most Basin. It was described from the Late Aquitanian-Early Burdigalian of the Mainz Basin and is additionally recorded for the Early Miocene of the Aquitaine Basin, the Hanau Basin, the Rhön Mountains, Southern Germany, and Moravia (Wenz 1923, Moayedpour 1977). Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake Infraorder Planorboinei Nordsieck, 1993 Superfamily Acroloxoidea Thiele, 1931 Family Acroloxidae Thiele, 1931 Subfamily Acroloxinae Thiele, 1931 Genus Acroloxus Beck, 1838 Type species. – Patella lacustris Linnaeus, 1758; subsequent designation by Herrmannsen (1846). Recent, Europe. Acroloxus decussatus (Reuss in Reuss & Meyer, 1849) Figure 3G, H 1849a Ancylus decussatus Rss. – Reuss in Reuss & Meyer, p. 11 (nomen nudum). *1849b A.[ncylus] decussatus m.; Reuss in Reuss & Meyer, p. 17, pl. 1, fig. 1. 1861 A.[ncylus] decussatus Reuss. – Reuss, p. 80. 1874 Ancylus (Velletia) decussatus Reuss. – Sandberger, p. 424, pl. 24, figs 2–2c. 1891 Ancylus (Velletia) decussatus Reuss. – Klika, p. 110, text-figs 109a–c. 1892 Ancylus (Velletia) decussatus Reuss. – Klika, p. 106, text-figs 109a–c. 1911 Ancylus (Velletia) decussatus Rss. – Kafka, p. 69. 1915 Velletia decussata (Reuss). – Fischer & Wenz, p. 57, unnumbered text-fig. on p. 58. ? 1915b Velletia decussata (Reuss). – Wenz, p. 43. 1916 Ancylus (Velletia) decussatus Reuss. – Thuma, p. 84. 1916 Acroloxus decussatus (Reuss). – Wenz, p. 181. 1917 Acroloxus decussatus (Reuss). – Wenz, p. 75. 1923 Acroloxus decussatus (Reuss). – Wenz, p. 1684 (pars). ? 1977 Acroloxus decussatus (Reuss). – Moayedpour, p. 62, pl. 3, figs 9, 10. Material. – One specimen (NHMW 1909/0001/0095). Dimensions. – Length: 2.5 mm, width: 1.8 mm, height: ca 0.8 mm. Description. – Apex raised, cap-like, inclined to the left; initial part of protoconch ca 200 μm in diameter, with smooth surface; surrounded by dense, collar-like pattern of radial and spiral threads; entire protoconch measuring ca 850 μm in maximum diameter; transition to teleoconch marked by distinct flattening of shell and almost abrupt cessation of radial-spiral pattern. Teleoconch regularly oval, covered by moderately distinct spiral growth lines. Discussion. – The clearly leftwards-inclined apex classifies this species as a true Acroloxus. It is the only Acroloxus species known from the Early Miocene of Europe. The Middle Miocene A. deperditolacustris (Gottschick, 1911) from Lake Steinheim can be distinguished by the rather ovoid shape with narrow posterior part. Acroloxus ucrainicus Gozhik & Prysjazhnjuk, 1978, from the late Sarmatian (Tortonian) of Mykolayiv, Ukraine, differs in the presence of several regularly- spaced spiral ridges on the teleoconch; the shape of the apex is not discernible from the illustration. Moayedpour (1977) follows the determinations by Fischer & Wenz (1914) and lists “A. decussatus” from Theobaldshof/Rhön, Germany. The illustrated specimen deviates slightly from ours in the higher-domed shell reminding of Ancylus and the higher apex. It remains unclear whether both specimens are conspecific. Occurrence. – Tuchořice, Korozluky, and Pyšná in the Most Basin. The records from the late Oligocene-Early Miocene of the Mainz and Hanau basins, the Burdigalian of the Rhön, the Aquitanian Öpfingen beds of Ehingen (Donau), and the Oligocene of Recollaine in Switzerland (Wenz 1923) are partly doubtful and need confirmation. Superfamily Planorboidea Rafinesque, 1815 Family Planorbidae Rafinesque, 1815 Subfamily Planorbinae Rafinesque, 1815 Genus Gyraulus Charpentier, 1837 Type species. – Planorbis albus Müller, 1774; subsequent designation by Dall (1870). Recent, Europe. Gyraulus dealbatus (Braun in Walchner, 1851) Figure 3I–N 1849a Planorbis applanatus Thom. – Reuss in Reuss & Meyer, p. 11 (non Planorbis applanatus Thomä, 1845). 1849a Planorbis exiguus Rss.; Reuss in Reuss & Meyer, p. 11 (nomen nudum). 1849b Pl.[anorbis] applanatus Thom. – Reuss in Reuss & Meyer, p. 38, pl. 4, fig. 8 (non Planorbis applanatus Thomä, 1845). 1849b Pl.[anorbis] exiguus m.; Reuss in Reuss & Meyer, p. 38, pl. 4, fig. 9. *1851 Planorbis dealbatus A. Braun; Braun in Walchner, p. 1134 (p. 50 in offprint), No. 264. 1861 Pl.[anorbis] cognatus Rss.; Reuss, p. 79, pl. 3, fig. 15. 1861 Pl.[anorbis] declivis A. Br. – Reuss, p. 79 (non Planorbis declivis Braun in Walchner, 1851 = Planorbis applanatus Thomä, 1845). 1861 Pl.[anorbis] exiguus Reuss. – Reuss, p. 80. ? 1869a Ptychospira deloplecta gen. et sp. nov.; Slavík, p. 261, pl. 4, figs 14, 15. 831 Bulletin of Geosciences • Vol. 89, 4, 2014 ? 1869b Ptychospira deloplecta gen. et sp. nov.; Slavík, p. 267, pl. 4, figs 14, 15. 1891 Planorbis cognatus Reuss. – Klika, p. 108, text-figs 106a–c. 1891 Planorbis declivis al. Braun. – Klika, p. 107, text-figs 104a, b (non Planorbis declivis Braun in Walchner, 1851 = Planorbis applanatus Thomä, 1845). 1891 Planorbis laevis Klein. – Klika, p. 109, text-figs 107a–c (non Planorbis laevis Klein, 1846 = Planorbis kleini Gottschick & Wenz, 1916). 1892 Planorbis cognatus Reuss. – Klika, p. 104, text-figs 106a–c. 1892 Planorbis declivis al. Braun. – Klika, p. 102, text-figs 104a, b (non Planorbis declivis Braun in Walchner, 1851 = Planorbis applanatus Thomä, 1845). 1892 Planorbis laevis Klein. – Klika, p. 104, text-figs 107a–c (non Planorbis laevis Klein, 1846 = Planorbis kleini Gottschick & Wenz, 1916). 1911 [Planorbis] declivis A. Br. – Kafka, p. 69 (non Planorbis declivis Braun in Walchner, 1851 = Planorbis applanatus Thomä, 1845). 1911 [Planorbis] laevis Kl. – Kafka, p. 69 (non Planorbis laevis Klein, 1846 = Planorbis kleini Gottschick & Wenz, 1916). 1916 [Planorbis] declivis Al. Braun. – Thuma, p. 84 (non Planorbis declivis Braun in Walchner, 1851 = Planorbis applanatus Thomä, 1845). 1916 [Planorbis] laevis v. Klein. – Thuma, p. 84. 1917 Gyraulus multiformis applanatus (Thomae). – Wenz, p. 75 (non Planorbis applanatus Thomä, 1845). 1923 Gyraulus (Gyraulus) trochiformis applanatus (Thomae). – Wenz, p. 1579 (only Most Basin records). 1923 Gyraulus (Gyraulus) trochiformis dealbatus (A. Braun). – Wenz, p. 1591 (cum syn.). ? 1923 deleplecta [sic] Slavik. – Wenz, p. 1111. ? 1925 Planorbis declivis Al. Braun for scalaris. – Petrbok, p. 3, unnumbered pl., figs 4, 6 (non Planorbis declivis Braun in Walchner, 1851 = Planorbis applanatus Thomä, 1845). 1964 Gyraulus trochiformis applanatus (Thomae, 1845). – Čtyroký et al., p. 138, fig. 4. 1977 Gyraulus (Gyraulus) trochiformis dealbatus (Braun). – Moayedpour, p. 60, pl. 2, figs 7–9. Material. – Several ten thousands of individuals (NHMW 1909/0001/0092, 2013/0572/0070–0072). Dimensions. – Diameter: 4.5 mm, height: 1.1 mm (Fig. 3I, J, N); diameter: 4.5 mm, height: 1.1 mm (Fig. 3K, M); diameter: 4.4 mm, height: 1.2 mm (Fig. 3L). Discussion. – Among the freshwater gastropods, Gyraulus dealbatus clearly dominates the assemblage concerning individual number. The morphological plasticity of this group of planorbids prompted earlier workers to split it into several species. Only three of these are today accepted, i.e. G. applanatus (including the synonym declivis), G. dealbatus (including the synonym cognatus and exiguus), and G. kleini (see Wenz 1923). After careful assessment of the rich material of the Vienna and Prague museums collections it was still not possible to detect and separate distinct morphological units. The differences mentioned and/or illustrated by several authors often correspond to different growth stages. In early ontogeny the shell is higher convex with less pronounced angle, the last whorl is distinctly larger in relation to the preceding whorls, and both umbilicus and apex seem deeper immersed. Because of allometric growth, whorl width outpaces whorl height, producing a flatter shell during ontogeny and forming a relatively wider umbilicus and apical concavity. Although some uncertainty remains, we regard all records of the Most Basin to refer to a single species. A determination as Gyraulus applanatus is clearly wrong as the typical pronounced basal keel (e.g. Gottschick 1920b) is absent in the studied material and was also never described or illustrated from Most Basin shells; they rather correspond to G. dealbatus. Petrbok (1925) described several irregularly scalariform shells, which he considered conspecific with the normally coiled Gyraulus from Tuchořice. Although we agree with Petrbok (1925), the poor preservation of the specimens, stored in the Prague collection, does not allow a clear identification. Comparable decoiled and scalariform shells were also described by Moayedpour (1977) from the Early Miocene of the Rhön Mountains, Germany, supporting Petrbok’s interpretation. Ptychospira deloplecta Slavík, 1869 was considered a juvenile Pupilloidea by Boettger (1870a) and Wenz (1923). Later, Petrbok (1925) identified the specimens as scalariform planorbids and suggested that they are conspecific with Gyraulus declivis (synonym of G. applanatus). Occurrence. – Known from Korozluky, Tuchořice, Lipno, and Pyšná in the Most Basin. Additionally recorded from the Early-Middle Miocene of the Aquitaine Basin, the Figure 3. A, F – Radix subovata (von Zieten, 1832), NHMW 2013/0572/0006. • B, C – Radix subovata (von Zieten, 1832), NHMW 2013/0572/0006. • D, E – Stagnicola subpalustris (Thomä, 1845), NHMW 2013/0572/0008. • G, H – Acroloxus decussatus (Reuss in Reuss & Meyer, 1849), NHMW 1909/0001/0095. • I, J, N – Gyraulus dealbatus (Braun in Walchner, 1851), NHMW 2013/0572/0070. • K, M – Gyraulus dealbatus (Braun in Walchner, 1851), NHMW 2013/0572/0071. • L – Gyraulus dealbatus (Braun in Walchner, 1851), NHMW 2013/0572/0072. 832 Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake B A C G F E H I L D J M K N 833 Bulletin of Geosciences • Vol. 89, 4, 2014 Mainz and Hanau basins, the Rhön Mountains, Southern Germany, Austria, Moravia, Hungary, Poland, Serbia, and Ukraine (Wenz 1923, Schlickum 1964, Čtyroký 1972, Moayedpour 1977, Gozhik & Prysjazhnjuk 1978, Piechocki 1997, Binder 2004, Kókay 2006, Prysjazhnjuk 2008). Many of these records need confirmation. to 50% of the preceding whorl. Additionally, there seems to be an angulation on both sides near the suture, producing a small funnel-shaped concavity. Hippeutis fasciatus Gottschick, 1920 from the Middle Miocene of Lake Steinheim is characterised by strong growth lines (see also Finger 1998). Moreover, the angulation is not central and the aperture not symmetrical in apertural view. Genus Hippeutis Charpentier, 1837 Occurrence. – Korozluky and Tuchořice in the Most Basin. It was also mentioned from the Aquitanian Öpfingen beds of Donaurieden, Baden-Württemberg (Wenz 1916). The record from the middle-late Burdigalian locality Trijebine in Serbia by Prysjazhnjuk (2008) can be only tentatively assigned to H. ungeri. The poor illustrations do not allow a clear classification. Type species. – Helix complanata Linnaeus, 1758; subsequent designation by Zilch in Wenz & Zilch (1959; see Welter-Schultes 2012 for discussion). Recent, Sweden. Hippeutis ungeri (Reuss in Reuss & Meyer, 1849) Figure 4A–C 1849a Pl.[anorbis] Ungeri Rss.; Reuss in Reuss & Meyer, p. 11 (nomen nudum). *1849b Pl.[anorbis] Ungeri m.; Reuss in Reuss & Meyer, p. 39, pl. 4, fig. 10. 1861 Pl.[anorbis] Ungeri Reuss. – Reuss, p. 79. 1874 Planorbis Ungeri Reuss. – Sandberger, p. 424, pl. 24, figs 1–1c. 1891 Planorbis ungeri Reuss. – Klika, p. 108, text-figs 105a, b. 1892 Planorbis ungeri Reuss. – Klika, p. 103, text-figs 105a, b. 1911 [Planorbis] Ungeri Rss. – Kafka, p. 69. 1916 [Planorbis] Ungeri Reuss. – Thuma, p. 84. 1916 Hippeutis ungeri (Reuss). – Wenz, p. 181. 1917 Hippeutis ungeri (Reuss). – Wenz, p. 75. 1923 Hippeutis (Hippeutis) ungeri (Reuss). – Wenz, p. 1649 (cum syn.). ? 2008 Hyppeutis (Hyppeutis) ungeri (Reuss, 1849). – Prysjazhnjuk, p. 88, pl. 1, figs 5, 6. Material. – One specimen (NHMW 1909/0001/0094), 6 specimens (NM-PM-P 768). Dimensions. – Diameter: 3.2 mm, height: 0.8 mm (Fig. 4A, C). Description. – Protoconch indiscernible, covered with sediment. Shell with distinct median angle, forming a weak but not offset keel. Aperture heart-shaped, with thin peristome. Moderately distinct, dense sigmoidal (in apertural view) growth lines cover the shell. Discussion. – Hippeutis subfontanus (Clessin, 1877) from the latest Burdigalian–early Langhian Silvana beds of Undorf, Hohenmemmingen, Mörsingen, and Zwiefaltendorf (Clessin 1877, Wenz 1923, Schlickum 1976) has a much stronger, blade-like keel and the last whorl is covering up 834 Genus Planorbarius Duméril, 1806 Type species. – Helix cornea Linnaeus, 1758; subsequent designation by Froriep (1806). Recent, Europe. Planorbarius cornu (Brongniart, 1810) Figure 4D–K *1810 Planorbis Cornu; Brongniart, p. 371, pl. 22, fig. 6. 1845 Planorbis solidus Nob. – Thomä, p. 153. 1849a Pl.[anorbis] pseudoammonius Voltz. – Reuss in Reuss & Meyer, p. 11 (non Helicites pseudammonius von Schlotheim, 1820). 1849a Pl.[anorbis] decussatus Rss.; Reuss in Reuss & Meyer, p. 11 (nomen nudum). 1849b Pl.[anorbis] pseudoammonius Voltz. – Reuss in Reuss & Meyer, p. 37, pl. 4, fig. 7 (non Helicites pseudammonius von Schlotheim, 1820). 1849b Pl.[anorbis] decussatus m.; Reuss in Reuss & Meyer, p. 39. 1849b Pl.[anorbis] excavatus Rss.; Reuss in Reuss & Meyer, pl. 4, fig. 11 (error pro decussatus). 1858 Planorbis solidus Thomæ. – Sandberger, p. 71, pl. 7, figs 8–8b. 1861 Pl.[anorbis] solidus Thom. – Reuss, p. 79. 1861 Pl.[anorbis] decussatus Reuss. – Reuss, p. 80. 1875 Planorbis cornu Brongniart. – Sandberger, p. 370 (pars). 1891 Planorbis cornu Brongniart. – Klika, p. 106, text-figs 103a, b. 1891 Planorbis cornu Brongniart. – Klika, p. 101, text-figs 103a, b. 1911 Planorbis cornu Brggt. – Kafka, p. 69. 1915 Planorbis cornu Brong. var. solida Tho. – Fischer & Wenz, p. 56. 1916 Planorbis cornu Brongniart. – Thuma, p. 84. 1917 Planorbis cornu Brongniart var. – Wenz, p. 74. Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake C D A B E F G H I J K Figure 4. A, C – Hippeutis ungeri (Reuss in Reuss & Meyer, 1849), NHMW 1909/0001/0094. • B – Hippeutis ungeri (Reuss in Reuss & Meyer, 1849), NM-PM-P 768. • D, F–G – Planorbarius cornu (Brongniart, 1810), NHMW 2013/0572/0010. • E, I, J – Planorbarius cornu (Brongniart, 1810), NHMW 2013/0572/0010. • H – Planorbarius cornu (Brongniart, 1810), NHMW 2013/0572/0073 (juvenile specimen). • K – Planorbarius cornu (Brongniart, 1810), NHMW 2013/0572/0074 (protoconch). 1923 Coretus cornu cornu (Brongniart). – Wenz, p. 1426. 1966 Planorbarius cornu (Brongniart). – Schlickum, p. 326, pl. 13, fig. 27. 1977 Planorbarius cornu cornu (Brongniart). – Moayedpour, p. 61, pl. 3, figs 7, 8. 2004 Planorbarius cornu (Brongniart, 1810). – Binder, p. 193, pl. 2, figs 2, 3. Material. – 3 specimens (NHMW 2013/0572/0010, 2013/0572/00073–0074), 429 specimens (NHMW 1909/0001/089). Dimensions. – Diameter: 18.0 mm, height: 7.2 mm (Fig. 4E, I, J); diameter: 20.5 mm, height: 8.0 mm (Fig. 4D, F, G). Discussion. – This species reminds of its (supposed) phylogenetic descendent, the Middle Miocene P. mantelli. Both species share the same general shape, the protoconch with spirally arranged pits, and the optional occurrence of more or less distinct spiral striae on the teleoconch (Harzhauser et al. 2014). However, adult shells of P. mantelli can be separated from those of P. cornu for being flatter with less bulbous whorls. 835 Bulletin of Geosciences • Vol. 89, 4, 2014 Wenz (1917) stated that the Bohemian shells have a rather flat body whorl in contrast to “typical” P. cornu from the Aquitanian–early Burdigalian of the Mainz Basin. However, this opinion cannot be followed herein as already the illustrations of Brongniart (1810) show a rather flat shell. Similarly, Thomä (1845) indicated a flattened “upper” (actually umbilical) part for specimens of Wiesbaden (Mainz Basin). Obviously unaware of the existence of P. cornu he introduced a new taxon for these shells, i.e. “Planorbis” solidus. It is considered a synonym of P. cornu by Wenz (1923) and this is followed herein. The record of the (Eocene!) species Planorbis pseudoammonius by Reuss in Reuss & Meyer (1849) is a misidentification and refers to P. cornu as well. Likewise, the “new species” Planorbis decussatus Reuss in Reuss & Meyer, 1849 represents juvenile individuals of P. cornu. Occurrence. – This taxon is wide-spread throughout the Late Oligocene and Early Miocene of Europe (Wenz 1923). Many records in the literature, especially those referring to Eocene or Middle Miocene localities, have to be checked. Known from Korozluky, Tuchořice, and Lipno in the Most Basin. Superorder Eupulmonata Haszprunar & Huber, 1990 Infraorder Acteophila Dall, 1885 Superfamily Ellobioidea H. & A. Adams, 1855 Family Carychiidae Jeffreys, 1830 Genus Carychiopsis Sandberger, 1872 sensu Strauch, 1977 Type species. – Pupa Dhorni Deshayes, 1863; subsequent designation by Tryon, 1884. Paleocene, France. Discussion. – The genus Carychiopsis was introduced by Sandberger (1872) without designating a type species. As he discussed two species (Pupa dhorni Deshayes, 1863 and P. alternans Deshayes, 1863) the principle of monotypy (ICZN Article 68.3) does not apply in contrast to the opinion of subsequent authors (e.g. Wenz & Zilch 1959). To our knowledge, the first author who listed Pupa dhorni as type species was Tryon (1884, p. 95). Although it is obvious that Tryon (1884) considered the single species listed with each genus as type, he did never explicitly define these as type species. Therefore, it may be disputable if this is in accordance with the ICZN. In this case the next author explicitly designating the type species of Carychiopsis was Cossmann (1889a, p. 340). No detailed analysis of the Paleocene Carychiopsis dhorni is available so far and therefore it remains unclear if the much younger Late Oligocene and Early Miocene spe836 cies treated by Strauch (1977) as Carychiopsis are really congeneric with the type species. Carychiopsis schwageri (Reuss, 1868) Figure 5A–D *1868 Pupa Schwageri sp. nov.; Reuss, p. 82, pl. 1, figs 5a–c. 1869a Carychium Schwageri Reuss sp. – Slavík, p. 265, pl. 4, figs 20, 21. 1869b Carychium Schwageri Reuss sp. – Slavík, p. 268, pl. 4, figs 20, 21. 1870a Carychium costulatum Sandb. – Boettger, p. 297. 1891 [Carychiopsis costulata Sandberger] var. schwageri Reuss. – Klika, p. 101, text-fig. 96. 1892 [Carychiopsis costulata Sandberger] var. schwageri Reuss. – Klika, p. 96, text-fig. 96. 1911 Carychiopsis costulata Sandb. var. Schwageri. – Kafka, p. 69. 1917 Carychiopsis costulata var. schwageri (Reuss). – Wenz, p. 73. 1923 Carychiopsis schwageri schwageri (Reuss). – Wenz, p. 1181. 1964 Carychiopsis schwageri schwageri (Reuss, 1868). – Čtyroký et al., p. 137, fig. 3. 1977 Carychium (Carychiopsis) schwageri (Reuss, 1868). – Strauch, p. 161 (pars), pl. 14, fig. 23, pl. 20, figs 7, 8. 1984 Carychium (Carychiopsis) schwageri (Reuss). – Prisyazhnyuk, p. 117, fig. 1a. 1995 Carychiopsis schwageri (Reuss, 1868). – Prisyazhnyuk & Stworzewicz, p. 268, fig. 1A. 1999 Carychiopsis schwageri (Reuss). – Esu, p. 332. non 1977 Carychium (Carychiopsis) schwageri (Reuss 1868). – Strauch, p. 161 (pars), pl. 20, figs 7–9 (= Carychium surai Stworzewicz, 1999a). non 1993 Carychium schwageri (Reuss, 1868). – Stworzewicz, p. 398 (= Carychiopsis prisyazhnyuki Stworzewicz, 1999a). Material. – 2 specimens (NHMW 2012/0572/0011), 159 specimens (NHMW 1909/0001/0082) (mixed with C. prisyazhnyuki Stworzewicz, 1999). Dimensions. – Diameter: 0.9 mm, height: 2.25 mm (Fig. 5A); diameter: 0.85 mm, height: 2.15 mm (Fig. 5B). Discussion. – Prisyazhnyuk (1984), Prisyazhnyuk & Stworzewicz (1995) and Stworzewicz (1999a) pointed out that two species from Tuchořice have been lumped as Carychiopsis schwageri. Of these, Carychiopsis schwageri is slender high-spired and has folded and wide columellar and parietal lamellae, whilst Carychiopsis prisyazhnyuki Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake Stworzewicz, 1999 is smaller, stout, low-spired and has simple lamellae. In addition, our SEM-investigations suggest that the protoconch of Carychiopsis schwageri is distinctly larger, higher and less convex (diameter: 260 μm, height: 180 μm) than in C. prisyazhnyuki, which attains about 200 μm in diameter and only 100 μm in height. Both species display a dense pattern of vaguely spirally arranged, tiny pits covering the entire protoconch. Due to the striking similarity in teleoconch morphology with near identical axial and spiral sculpture and the identical protoconch microsculpture, we refrain from placing both species in different (sub)genera as proposed by Prisyazhnyuk & Stworzewicz (1995). The main arguments for that were the “Saraphia-like” columellar and parietal lamellae in C. schwageri. The width and degree of undulation of the lamellae, however, are quite variable in our material of C. schwageri. Occurrence. – This species is known only from Tuchořice, Pyšná and the Kralupy drilling; occurrences from the Middle Miocene of Opole (Poland) mentioned by Strauch (1977) represent Carychium surai Stworzewicz, 1999. Carychiopsis prisyazhnyuki Stworzewicz, 1999 Figure 5E–G 1977 Carychiopsis schwageri (Reuss). – Moayedpour, p. 58, pl. 2, figs 4, 5 (non Pupa Schwageri Reuss, 1868). 1984 Carychium (Carychiopsis) sp. – Prisyazhnyuk, p. 117, fig. 1b. 1995 Carychiopsis sp. nov. – Prisyazhnyuk & Stworzewicz, p. 268, fig. 1B. *1999a Carychiopsis prisyazhnyuki sp. nov.; Stworzewicz, p. 262, figs 1–4 (cum syn.). Material. – 3 specimens (NHMW 2012/0572/0012). Dimensions. – Diameter: 0.9 mm, height: 1.9 mm (Fig. 5E, F). Discussion. – See C. schwageri. Prisyazhnyuk (1984) was the first who described this rather stout species as Carychium (Carychiopsis) sp. and later Stworzewicz (1999a) introduced prisyazhnyuki as name for this species, which is based on a holotype from Bełchatów (Poland) and numerous paratypes including one specimen from Tuchořice. Therefore, parts of the references listed with C. schwageri may refer also to Carychium prisyazhnyuki Stworzewicz, 1999. Occurrence. – Known from Tuchořice and the middle to late Burdigalian locality Bełchatów (Bel-C). The stout outline and low protoconch of the shells from Theobalds- hof/Rhön in Germany, described by Moayedpour (1977) as C. schwageri, suggest that they represent C. prisyazhnyuki Stworzewicz, 1999. Genus Carychiella Strauch, 1977 Type species. – Carychium eumicrum Bourguignat, 1857; by original designation. Miocene, Europe. Carychiella eumicrum (Bourguignat, 1857) Figure 5H–J *1857 Carychium eumicrum Bourguignat, p. 223. 1860 Carychium eumicrum Bourguignat. – Bourguignat, p. 53, pl. 11, figs 3, 4. 1869a Carychium nanum Sandb. – Slavík, p. 266, pl. 4, figs 22, 23. 1869b Carychium nanum Sandb. – Slavík, p. 269, pl. 4, figs 22, 23. 1870a [Carychium nanum] Varietät major m. – Boettger, p. 297, pl. 13, figs 8a, b. 1891 [Carychium minutissimum Al. Br.] Böttgeri. – Flach, p. 58. 1891 Car.[ychium] majus Böttg. – Flach, p. 58. 1891 Carychium maius Boettger. – Klika, p. 102, text-figs 98a, b. 1891 [Carychium minutissimum Al. Braun] var. boettgeri Flach. – Klika, p. 102, text-figs 97a, b. 1892 Carychium maius Boettger. – Klika, p. 98, text-figs 98a, b. 1892 [Carychium minutissimum Al. Braun] var. boettgeri Flach. – Klika, p. 97, text-figs 97a, b. 1911 Carychium minutissimum Al. Br. var. Boettgeri Fl. – Kafka, p. 69. 1911 Carychium maius Bttg. – Kafka, p. 69. 1917 Carychium nanum boettgeri Flach. – Wenz, p. 73. 1923 Carychium eumicron eumicron [sic] Bourguignat. – Wenz, p. 1187. 1923 Carychium eumicron boettgeri Flach. – Wenz, p. 1189. 1923 Carychium majus Boettger. – Wenz, p. 1191. 1964 Carychium cf. majus Boettger, 1870. – Čtyroký et al., table for p. 149. 1977 Carychium (Carychiella) eumicron [sic] Bourguignat, 1857 s.l. – Strauch, p. 159, pl. 14, figs 13–15, pl. 17, fig. 53, pl. 19, fig. 76. 1995 Carychium eumicrum Bourguignat, 1860. – Prisyazhnyuk & Stworzewicz, p. 270. 1999a Carychium eumicrum Bourguignat. – Stworzewicz, p. 264, figs 5, 6 (cum syn.). 2006 Carychium (Carychiella) eumicron eumicron [sic] Bourguignat, 1857. – Kókay, p. 48, pl. 16, fig. 5. 2013 Carychium (Carychiella) eumicron [sic] Bourguignat, 1857. – Salvador, p. 2, fig. 1. 837 Bulletin of Geosciences • Vol. 89, 4, 2014 Material. – 3 specimens (NHMW 2013/0572/0013), >13,200 specimens (NHMW 1909/0001/0083, 1909/0001/0084). Dimensions. – Diameter: 0.55 mm, height: 1.05 mm (Fig. 5J); diameter: 0.5 mm, height: 0.78 mm (Fig. 5I); diameter: 0.55 mm, height: 1.05 mm (Fig. 5H). Discussion. – This is the smallest and by far most abundant species at Tuchořice (based on the NHMW collections). It is the type species of Carychiella Strauch, 1977, which was defined by Strauch (1977) based on the minute size, smooth shell, simple columellar and parietal lamellae and characteristic aperture with only three denticles. In addition, the punctate microsculpture of the proto- and teleoconch, as revealed by SEM studies on herein presented material and on Middle Miocene shells from the Rein Basin (Harzhauser et al. 2014), might be characteristic for this genus. Occurrence. – Carychium eumicrum appears during the Late Oligocene (Hochheim, Germany) and is recorded from the Early Miocene of Tuchořice, Lipno and Kralupy drilling (Czech Republic) and Early/Middle Miocene of Undorf and Sandelzhausen (S Germany) and from the Middle Miocene of Poland (Bełchatów, Opole), Austria (Rein Basin) and Hungary (Stworzewicz 1999a, Kókay 2006, Salvador 2013, Harzhauser et al. 2014). Infraorder Stylommatophora Schmidt, 1855 Clade Orthuretha Pilsbry, 1900b Superfamily Cochlicopoidea Pilsbry, 1900b (1879) Family Azecidae Watson, 1920 Genus Azeca Fleming, 1828 Type species. – Turbo tridens Pulteney, 1799 [= Azeca goodalli (Férussac, 1821)]. Recent, Europe. Azeca pumila Slavík, 1869 Figure 5K–M *1869a Azeca pumlia sp. nov.; Slavík, p. 252, pl. 4, figs 18, 19. 1869b Azeca pumlia sp. nov.; Slavík, p. 264, pl. 4, figs 18, 19. 1891 Azeca pumila Slavík. – Klika, p. 73, text-figs 70a, b. 1892 Azeca pumila Slavík. – Klika, p. 70, text-figs 70a, b. 1911 Azeca pumila Sl. – Kafka, p. 68. 1917 Azeca pumila Slavik. – Wenz, p. 71. 1923 Azeca (Azeca) monocraspedon pumila Slavik. – Wenz, p. 1095 (cum syn.). 1925 Azeca danesi sp. nov.; Petrbok, p. 2, unnumbered pl., fig. 2. 1925 Azeca pumila Slavík. – Petrbok, p. 2, unnumbered pl., fig. 1. Material. – 2 specimens (NHMW 2013/0572/0014), 6 specimens (NHMW 1909/0001/0056, 1909/0001/0060, 1909/0001/0062), 1 specimen (NM-PM-P 440, holotype of Azeca danesi Petrbok, 1925). Dimensions. – Diameter: 1.6 mm, height: 3.7 mm (Fig. 5K); diameter: 1.6 mm, height: 3.4 mm (Fig. 5L); diameter: 1.7 mm, height: 3.9 mm (= Azeca danesi Petrbok, 1925, Fig. 5M). Discussion. – The status of Azeca pumila Slavík, 1869, and A. monocraspedon Slavík, 1869 as distinct species was doubted by Boettger (1870a) and Wenz (1917, 1923). Nevertheless, both species differ considerably in the presence of well-developed columellar and palatal denticles in A. pumila. These are absent in A. monocraspedon, which has a broad and low columellar swelling, a narrow but strongly protruding parietal denticle and completely smooth outer lip. Azeca danesi Petrbok, 1925 is based on a single specimen, which is re-illustrated herein. It differs from Azeca pumila only in the absence of the parietal denticle and the slightly swollen parietal area. The re-investigation of the specimen, however, showed that parts of the columellar and parietal callus are broken off. This element forms a thin sheet in A. pumila bearing the parietal denticle. Therefore, A. danesi is most probably only an incompletely preserved Azeca pumila. Occurrence. – Only known from Tuchořice; a further occurrence was mentioned from the Middle Miocene of Opole by Andreae (1902b) without illustration and sufficient description. Figure 5. A–C – Carychiopsis schwageri (Reuss, 1868), NHMW 2013/0572/0011 (C – detail of the microsculpture). • D – Carychiopsis schwageri (Reuss, 1868) (protoconch of A–C). • E, F – Carychiopsis prisyazhnyuki Stworzewicz, 1999, NHMW 2013/0572/0012. • G – Carychiopsis prisyazhnyuki Stworzewicz, 1999 (protoconch of E, F). • H–J – Carychiella eumicrum (Bourguignat, 1857), NHMW 2013/0572/0013. • K, L – Azeca pumila Slavík, 1869, NHMW 2013/0572/0014. • M – Azeca pumila Slavík, 1869, NM-PM-P 440 (holotype of Azeca danesi Petrbok, 1925). • N, O – Azeca monocraspedon Slavík, 1869, NHMW 2013/0572/0015. • P–R – Hypnophila subrimata (Reuss in Reuss & Meyer, 1849), NHMW 2013/0572/0016. • S, T – Cochlicopa dormitzeri (Reuss in Reuss & Meyer, 1849), NHMW 2013/0572/0017. 838 Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake A H N D B F E I J O P G C L K Q M T S R 839 Bulletin of Geosciences • Vol. 89, 4, 2014 Azeca monocraspedon Slavík, 1869 Figure 5N–O *1869a Azeca monocraspedon sp. nov.; Slavík, p. 252, pl. 4, figs 16, 17. 1869b Azeca monocraspedon sp. nov.; Slavík, p. 263, pl. 4, figs 16, 17. 1875 Azeca monocraspedon Slavíc [sic]. – Sandberger, p. 434, pl. 24, figs 20–20b. 1891 Azeca monocraspedon Slavík. – Klika, p. 74, text-figs 71a, b. 1892 Azeca monocraspedon Slavík. – Klika, p. 71, text-figs 71a, b. 1911 [Azeca] monocraspedon Sl. – Kafka, p. 68. 1923 Azeca (Azeca) monocraspedon monocraspedon Slavík. – Wenz, p. 1094 (cum syn.). 1925 Azeca monocraspedon Slavík. – Petrbok, p. 2, unnumbered pl., fig. 3. Material. – One specimen (NHMW 2013/0572/0015), 2 specimens (NHMW 1909/0001/0061). Dimensions. – Diameter: 1.45 mm, height: 3.5 mm (Fig. 5N); diameter: 1.5 mm, height: 3.5 mm (Fig. 5O). Discussion. – See above. Occurrence. – Only known from Tuchořice. Genus Hypnophila Bourguignat, 1858 Type species. – Bulimus pupaeformis Cantraine, 1835; subsequent designation by Letourneux & Bourguignat, 1887. Recent, S Europe, N Africa. Hypnophila subrimata (Reuss in Reuss & Meyer, 1849) comb. nov. Figure 5P–R 1849a Achatina subrimosa Rss.; Reuss in Reuss & Meyer, p. 11 (nomen nudum). *1849b A.[chatina] subrimata m.; Reuss in Reuss & Meyer, p. 31, pl. 3, fig. 9. 1861 Gl.[andina] lubricella A. Br. sp. – Reuss, p. 70 (non Bulimus lubricus var. lubricella Porro, 1838). 1891 [Cionella lubricella A. Braun] var. subrimata Reuss. – Klika, p. 73, text-fig. 69. 1892 [Cionella lubricella A. Braun] var. subrimata Reuss. – Klika, p. 70, text-fig. 69. 1897 Cionella splendens A. Braun. – Babor, p. 17 (non Achatina splendens Braun in Walchner, 1851). 1911 [Cionella] lubricella A. Br. – Kafka, p. 68 (non Bulimus lubricus var. lubricella Porro, 1838). 840 1911 [Cionella lubricella A. Br.] var. subrimata Rss. – Kafka, p. 68. 1914 Cochlicopa subrimata (Reuss). – Wenz, p. 111, pl. 8, fig. 35. 1916 Cochlicopa subrimata (Reuss). – Wenz, p. 177. 1917 Cochlicopa subrimata (Reuss). – Wenz, p. 72. 1923 Cochlicopa subrimata subrimata (Reuss). – Wenz, p. 1102 (pars). non 1977 Cochlicopa subrimata (Reuss). – Moayedpour, p. 62, pl. 3, figs 11, 12. non 1998 Cochlicopa subrimata (Reuss, 1852). – Finger, p. 18, pl. 12, fig. F. Material. – 2 specimens (NHMW 2013/0572/0016), 58 specimens (NHMW 1909/0001/0058, 1909/0001/0059), 1 specimen (NM-PM-P 451). Dimensions. – Diameter: 2.3 mm, height: 5.3 mm (Fig. 5P); diameter: 2.1 mm, height: 5.1 mm (Fig. 5Q). Discussion. – This species was traditionally treated as Cochlicopa Férussac, 1821. The stout pupoid shell with nearly flat whorls and weakly incised sutures, however, suggest a relation with the Azecidae. Consequently, we place it in Hypnophila based on the simple aperture. The species differs clearly from the more slender Cochlicopa dormitzeri (Reuss in Reuss & Meyer, 1849), which has a much more elongate aperture and more convex whorls. For the first time, SEM studies of H. subrimata document the presence of densely spaced spiral grooves and of a delicate spiral thread along the upper suture of the first teleoconch whorl. Both elements fade out within the second teleoconch whorl and the rest of the shell is smooth aside from faint growth lines. Occurrence. – Known from Tuchořice, Lipno and Korozluky. The shell fragment from the Middle Miocene of Steinheim (Germany) identified as Cochlicopa subrimata by Finger (1998) is clearly not conspecific due to its conical spire and the convex whorls. Similarly, the specimen from Theobaldshof/Rhön illustrated by Moayedpour (1977) differs clearly in its convex whorls and the deeply concave columella. Therefore, the numerous Oligocene and early Miocene records listed by Wenz (1923) should be treated with caution. Family Cochlicopidae Pilsbry, 1900b Genus Cochlicopa Férussac, 1821 Type species. – Helix lubrica Müller, 1774; subsequent designation by Westerlund, 1902. Recent, Europe. Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake Cochlicopa dormitzeri (Reuss in Reuss & Meyer, 1849) Figure 5S–T 1849a Achatina Dormitzeri Rss.; Reuss in Reuss & Meyer, p. 12 (nomen nudum). *1849b A.[chatina] Dormitzeri m.; Reuss in Reuss & Meyer, p. 31, pl. 3, fig. 10. 1861 Gl.[andina] Dormitzeri Reuss. – Reuss, p. 70. 1891 Cionella Dormitzeri Reuss. – Klika, p. 71, text-fig. 68a, b. 1891 Cionella dormitzeri Reuss. – Thuma, p. 83. 1892 Cionella Dormitzeri Reuss. – Klika, p. 69, text-fig. 68a, b. 1911 Cionella Dormitzeri Rss. – Kafka, p. 68. 1923 Cochlicopa subrimata dormitzeri (Reuss). – Wenz, p. 1106. Material. – 2 specimens (NHMW 2013/0572/0017), 3 specimens (NHMW 1909/0001/0057). Dimensions. – Diameter: 1.7 mm, height: 4.5 mm (Fig. 5S); diameter: 1.7 mm, height: 4.3 mm (Fig. 5T). Discussion. – This rare species is recognised easily by its slender shell, high spire, slightly convex whorls and high, elongate aperture. In contrast to all Azecidae and Cochlicopidae of the Most Basin, it has a comparatively high and convex protoconch. 1861 B.[ulimus] complanatus Reuss. – Reuss, p. 69. 1875 Bulimus (Petraeus) complanatus Reuss. – Sandberger, p. 433, pl. 24, figs 11, 11a. 1891 Buliminus (Medaea?) complanatus Reuss. – Klika, p. 69, text-figs 64a, b. 1891 Buliminus (Petraeus) turgidulus (Sandberger). – Klika, p. 69, text-figs 65a, b [non Bulimus (Petraeus) turdigulus Sandberger, 1875]. 1892 Buliminus (Medaea?) complanatus Reuss. – Klika, p. 66, text-figs 64a, b. 1892 Buliminus (Petraeus) turgidulus (Sandberger). – Klika, p. 67, text-figs 65a, b [non Bulimus (Petraeus) turdigulus Sandberger, 1875]. 1916 Buliminus (Medaea?) complanatus Reuss. – Thuma, p. 83. 1917 Buliminus (?Napaeus) complanatus (Reuss). – Wenz, p. 64. 1923 Ena (Napaeus) complanata (Reuss). – Wenz, p. 1074. ? 2004 Napaeus cf. complanatus (Reuss, 1849). – Binder, p. 196, pl. 5, figs 2a, b. non 1911 Bulimus (Petræus) complanatus Reuss. – Gaál, p. 70, pl. 2, figs 9, 12 (= Ena gaali Wenz, 1919). non 2006 Napaeus complanatus (Reuss), 1849. – Kókay, p. 73, pl. 27, figs 10–12. Material. – 2 specimens (NHMW 2013/0572/0018), 61 specimens (NHMW 1909/0001/0055). Occurrence. – Only known from Korozluky and Tuchořice. Dimensions. – Diameter: 8.5 mm, height: 17.9 mm (Fig. 6A, B); diameter: 8.4 mm, height: 16.9 mm (Fig. 6C, D). Superfamily Enoidea Woodward, 1903 Family Enidae Woodward, 1903 Description. – A stout ovate-conic shell with weakly cyrtoconoid spire and weakly convex whorls and moderately dome-shaped protoconch. The glossy shell bears densely spaced, strongly prosocline growth lines with a narrow and shallow prosocyrt sulcus close to the weakly incised upper suture. Broadly flared outer and columellar lips are connected by a moderately developed parietal callus. The reflected columellar lip delimitates a narrow and moderately deep umbilical chink. A delicate central swelling appears at the transition from the wide inner lip into the thin and straight columella. The transition from the convex protoconch whorl to the weakly convex early teleoconch is much less abrupt than in most extant Mastus and Ena species, which often display somewhat bulbous protoconchs. Genus Mastus Beck, 1837 Type species. – Helix pupa Linnaeus, 1758; subsequent designation by Herrmannsen (1846). Recent, S Europe, N Africa. Mastus complanatus (Reuss in Reuss & Meyer, 1849) Figure 6A–D 1849a Bulimus complanatus Rss.; Reuss in Reuss & Meyer, p. 11 (nomen nudum). 1849a Bulimus Meyeri Rss.; Reuss in Reuss & Meyer, p. 11 (nomen nudum). *1849b Bulimus complanatus m.; Reuss in Reuss & Meyer, p. 29, pl. 3, fig. 4. 1849b B.[ulimus] Meyeri m.; Reuss in Reuss & Meyer, p. 29, pl. 3, fig. 5. 1855 Bulimus complanatus Reuss. – Pictet, p. 27, pl. 57, fig. 14. Discussion. – Since Wenz (1917) this species has been assigned to Napaeus Albers, 1850, which is a genus endemic to the Canary Islands (Henriquez et al. 1993). Although Wenz (1923) considered most Eocene to Miocene Eninae as Napaeus, it seems doubtful that Central and SouthEastern European fossil species are indeed congeneric with this genus. Aside from this palaeogeographic argument, the 841 Bulletin of Geosciences • Vol. 89, 4, 2014 Bohemian species lacks the prominent microsculpture of the early whorls of Napaeus as documented by Henriquez et al. (1993) for six different species. In addition, the protoconchs of these species are distinctly more erect. As already discussed by Klika (1891) the shell outline of the Bohemian species is highly reminiscent of the extant Ena raddei (Kobelt, 1880) from the Caucasus region, which is the type species of the subgenus Caucasicola Hesse, 1917. This species lacks the reflected peristome of the Bohemian species and displays a delicate radial sculpture on early spire whorls. Therefore, we refrain from assigning the Miocene species to Caucasicola. According to Eike Neubert (pers. comm.) this species does not belong to the genus Ena Turton, 1831. In his online checklist on the genera of fossil land snails (Stylommatophora) of western and central Europe, Hartmut Nordsieck (http://www.hnords.de/) lists this species as Mastus, which is provisionally followed herein. As pointed out by Nordsieck (written comm. 2014) this genus is represented during the Miocene by several similar species in the Caucasus region described by Steklov (1966). The specimen described as Buliminus turgidulus by Klika (1891, 1892) is a poorly preserved internal mould and is most probably conspecific with Mastus complanatus. Occurrence. – Known from Korozluky, Tuchořice, Lipno and Pyšná. This species was also described from the Ottnangian of Oberdorf in Austria (Binder 2004). The status of these poorly preserved specimens is difficult to evaluate. The occurrence from the Lower Miocene of the Somlóvásárhely drilling in Hungary (Kókay 2006) is most probably not conspecific as the subsutural furrow below the suture, as described by Kókay (2006), does not occur in the Bohemian species. Mastus filocinctus (Reuss, 1861) Figure 6E–G *1861 B.[ulimus] filocinctus Reuss; Reuss, p. 69, pl. 2, fig. 5. 1875 Bulimus (Petraeus) filocinctus Reuss. – Sandberger, p. 433, pl. 24, figs 12–12b. 1891 Buliminus (Petraeus) filocinctus Reuss. – Klika, p. 68, text-figs 63/1, 2a, b. 1892 Buliminus (Petraeus) filocinctus Reuss. – Klika, p. 65, text-figs 63/1, 2a, b. 1911 Buliminus (Petraeus) filocinctus Rss. – Kafka, p. 68. 1917 Buliminus (Napaeus) filocinctus (Reuss). – Wenz, p. 64. 1923 Ena (Napaeus) filocincta (Reuss). – Wenz, p. 1075 (cum syn.). Material. – One specimen (NHMW 2013/0572/0019), one specimen (NHMW 1909/0001/0054), 4 specimens (NM-PM-P 751–754). 842 Dimensions. – Diameter: 4.8 mm, height: 13.1 mm (Fig. 6E, NHMW 2013/0572/0019), diameter: 5.2 mm, height 15 mm (Fig. 6F, NM-PM-P 753), diameter: 5.5 mm, height 13 mm (Fig. 6G, NM-PM-P 752). Discussion. – A rare species, which is characterised by a slender shell, a dome-shaped protoconch, densely spaced, prominent, prosocline growth lines, a thickened and reflected, ovate and moderately wide peristome, connected by a thickened parietal callus and a conspicuous spiral thread along the upper suture. The seemingly canaliculate suture between the third and fourth spire whorls of specimen NHMW 2013/0572/0019 is an artefact from incorrect gluing. This species is rather variable concerning slenderness. Occurrence. – Only known from Tuchořice. Superfamily Pupilloidea Turton, 1831 Family Pupillidae Turton, 1831 Genus Paracoryna Flach, 1891 Type species. – Pupa (Coryna) Diezi Flach, 1891; subsequent designation by Pilsbry (1924). Early Miocene, Czech Republic. Paracoryna diezi (Flach, 1891) Figure 6H–J *1891 Pupa (Coryna Westl.) Diezi n. form; Flach, p. 49, pl. 3, figs 1a, b. 1891 Coryna diezi Flach. – Klika, p. 94, text-figs 89a, b. 1892 Coryna diezi Flach. – Klika, p. 90, text-figs 89a, b. 1911 Coryna diezi Flach. – Kafka, p. 69. 1917 Agardhia diezi (Flach). – Wenz, p. 66. 1923 Agardhia diezi (Flach). – Wenz, p. 1037 (cum syn.). 1924 Agardhia (Paracoryna) diezi. – Pilsbry, p. 133, pl. 19, figs 17, 18. Material. – 2 specimens (NHMW 2013/0572/0020), 6 specimens (NHMW 1909/0001/0075). Dimensions. – Diameter: 1.15 mm, height: 1.75 mm (Fig. 6H); diameter: 1.1 mm, height: 1.85 mm (Fig. 6I). Description. – Stout pupoid shell consisting of 3–4 strongly convex whorls, separated by deeply incised sutures, and a moderately convex protoconch whorl with coarsely malleated surface. Teleoconch covered by densely spaced, prosocline, wrinkle-like and discontinuous axial ribs. Narrow umbilicus; wide aperture with thin peristome, slightly widened lips; no denticles or lamellae are developed. Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake Discussion. – This species is rather rare and our shells do not fully agree with the original description of Flach (1891), who presents a shell with an additional whorl, a higher last whorl and higher aperture. The size relation suggests that our shells might rather be subadult specimens. This species was placed by several authors in Coryna Westerlund, 1887 (non Bosc, 1802) and Agardhia Gude, 1911 (pro Coryna Westerlund), which are synonyms of Argna Cossmann, 1889b according to Aellen & Finet (1990). Even Pilsbry (1924) placed it under the subgenus Paracoryna Flach within Agardhia. The protoconch morphology and sculpture of our specimens differ considerably from that of Miocene Argna species, such as A. oppoliensis (Andreae, 1902) with depressed domical and smooth protoconch (see Stworzewicz 1999b and Harzhauser & Binder 2004). Comparable protoconch structures, however, are present in several Miocene Pupillidae (see Finger 1998, Harzhauser et al. 2008). Occurrence. – Only known from Tuchořice. Family Strobilopsidae Wenz, 1915 2006 Strobilops fischeri Wenz, 1915. – Kókay, p. 71, pl. 26, figs 5, 6. Material. – 2 specimens (NHMW 2013/0572/0021), 42 specimens (NHMW 1909/0001/0021). Dimensions. – Diameter: 2.0 mm, height: 1.15 mm; diameter: 2.3 mm, height: 1.3 mm. Discussion. – This species was described in great detail by Wenz in Fischer & Wenz (1914) and Wenz (1915a). It is characterised by a smooth protoconch, a low spire and its sculpture consists of low and somewhat irregular axial ribs, often separated by weaker and discontinuous secondary ribs; the umbilicus is narrow but deep. The aperture reveals two prominent and thin parietal lamellae with weak knots, two short basal folds and a weak and narrow columellar fold. Aside from the depressed outline, its sculpture and apertural features correspond fully to Eostrobilops as defined by Pilsbry (1927). Type species. – Strobilops hirasei Pilsbry, 1908b; original designation. Recent, Korea. Occurrence. – Known from Tuchořice and Korozluky. Additional occurrences are documented from the Early Miocene of Hungary (Somlóvásárhely drilling, Kókay 2006), Theobaldshof/Rhön in Germany (Moayedpour 1977) and Bełchatów in Poland (Stworzewicz 1999b). The youngest records are mentioned from the Middle Miocene of Bełchatów-B by Stworzewicz (1999b). Strobilops (Eostrobilops) fischeri (Wenz in Fischer & Wenz, 1914) Figure 6R–W Strobilops (Eostrobilops) elasmodonta (Reuss, 1861) Figure 6K–M Genus Strobilops Pilsbry, 1893 Subgenus Eostrobilops Pilsbry, 1927 1891 Strobilus diptyx Boettger. – Klika, p. 34, text-figs 26a–c (non Helix diptyx Boettger, 1870b). 1892 Strobilus diptyx Boettger. – Klika, p. 33, text-figs 26a–c (non Helix diptyx Boettger, 1870b). 1911 [Strobilus] diptyx Bttg. – Kafka, p. 67 (non Helix diptyx Boettger, 1870b). *1914 Strobilops fischeri sp. nov.; Wenz in Fischer & Wenz, p. 107. 1915a Strobilops (Strobilops) fischeri sp. nov. – Wenz, p. 78, fig. 6, pl. 4, fig. 5. 1917 Strobilops (Strobilops) fischeri Wenz. – Wenz, p. 70. 1916 [Strobilus] diptix Boettger. – Thuma, p. 83 (non Helix diptyx Boettger, 1870b). 1923 Strobilops (Strobilops) fischeri Wenz. – Wenz, p. 1047. 1927 Strobilops fischeri Wenz. – Pilsbry, p. 6. 1977 Strobilops (Strobilops) fischeri Wenz. – Moayedpour, p. 64, pl. 4, figs 10–12, pl. 5, figs 1, 2. 1999 Eostrobilops fischeri Wenz. – Esu, p. 332. 1999b Strobilops fischeri Wenz, 1914. – Stworzewicz, p. 157, figs 48–50. *1861 H.[elix] elasmodonta Reuss; Reuss, p. 66, pl. 1, fig. 6. 1870a Hyalinia elasmodonta Reuss. – Boettger, p. 287. 1875 Strobilus elasmodonta Reuss. – Sandberger, p. 442, pl. 24, figs 24–24b. 1891 Strobilus elasmodonta Reuss. – Klika, p. 33, text-figs 24a, 25b, c (non 25a = Discostrobilops uniplicatus). 1892 Strobilus elasmodonta Reuss. – Klika, p. 33, text-figs 24a, 25a–c. 1911 [Strobilus] elasmodonta Rss. – Kafka, p. 67. 1912 Str.[obilus] elasmodonta (Reuss). – Jooss, p. 35. 1915a Strobilops (Str.[obilops]) elasmodonta (Reuss). – Wenz, p. 77, pl. 4, figs 7a–c. 1916 Strobilus elasmodonta Reuss. – Thuma, p. 83. 1917 Strobilops (Strobilops) elasmodonta (Reuss). – Wenz, p. 70. 1923 Strobilops (Strobilops) elasmodonta (Reuss). – Wenz, p. 1046. 1927 Strobilops elasmodonta (Reuss). – Pilsbry, p. 6. 1964 Strobilops elasmodonta (Reuss, 1860). – Čtyroký et al., table for p. 149. 843 Bulletin of Geosciences • Vol. 89, 4, 2014 1970a St.[robilops] elasmodonta (Reuss). – Schlickum, p. 83. 1977 Strobilops (Strobilops) cf. elasmodonta (Reuss). – Moayedpour, p. 64, pl. 5, figs 3, 4. non 2003 Strobilus elasmodonta Reuss. – Mikuláš et al., p. 94. Material. – 2 specimens (NHMW 2013/0572/0022), 57 specimens (NHMW 1909/0001/0020). Dimensions. – Diameter: 2.1 mm, height: 1.3 mm (Fig. 6K); diameter: 2.1 mm, height: 1.35 mm (Fig. 6L). Discussion. – This species is readily distinguished from the syntopic Strobilops (Eostrobilops) fischeri by its higher, conical spire and narrower umbilicus. In contrast to S. fischeri, its protoconch is weakly granulose on the initial part; these granules grade into faint, axially arranged wrinkles towards the teleoconch, which bears densely spaced and low axial ribs. Pilsbry (1927) discussed S. (E.) elasmodonta to belong probably to the Strobilops diptyx-fischeri-boettgeri-subconoidea-duvali lineage, which would imply a position within Eostrobilops after Pilsbry (1927). Doubts remained because of the presence of an interparietal lamella in S. (E.) elasmodonta. Similarly, Schlickum (1970) doubted that S. (E.) elasmodonta belongs to Eostrobilops although he kept his newly described “elasmodonta-like” Strobilops senckenbergi within Eostrobilops. In contrast to the statement of Schlickum (1970a), the Bohemian species has a narrow umbilicus, which may be covered by the reflected lip in some specimens. Therefore, only the weak interparietal fold remains as problematic feature. Herein, we provisionally assign S. (E.) elasmodonta to the subgenus Eostrobilops based on its general shell shape, reduced sculpture and narrow umbilicus. Occurrence. – Known from Korozluky, Tuchořice, Lipno and the Chomutov drillings in the Most Basin. The occurrence from the Lower Oligocene of Dětaň (Czech Republic), mentioned by Mikuláš et al. (2003), needs verification but seems unlikely in respect to the age difference. A further occurrence from the Burdigalian of Theobaldshof/Rhön (Germany), described by Moayedpour (1977), is likely based on the characteristic outline although the preservation excludes a clear identification. Subgenus Discostrobilops Pilsbry, 1927 Type species. – Helix Hubbardi Brown, 1861; original designation. Recent, Mexico and southern USA. Strobilops (Discostrobilops) uniplicatus (Braun in Walchner, 1851) Figure 6N–Q *1851 Helix uniplicata A. Braun; Braun in Walchner, p. 1138, p. 54 in offprint, No. 320. 1858 Helix uniplicata Al. Braun. – Sandberger, p. 35, pl. 3, figs 7–7c. 1861 H.[elix] uniplicata A. Br. – Reuss, p. 68. 1875 Strobilus uniplicatus A. Braun. – Sandberger, p. 406, pl. 23, figs 24–24b. 1891 Strobilus uniplicatus Al. Braun. – Klika, p. 32, text-figs 24b, c, 25a (non 24a = Eostrobilops elasmodonta). 1892 Strobilus uniplicatus Al. Braun. – Klika, p. 31, text-figs 24a–c. 1911 Strobilus uniplicatus A. Br. – Kafka, p. 67. 1914 Strobilops uniplicata (Sdbg.). – Wenz in Fischer & Wenz, p. 106, pl. 7, fig. 33. 1915a Strobilops (Str.[obilops]) uniplicata (Sandberger). – Wenz, p. 76, pl. 4, figs 8a–c. 1916 Strobilops uniplicata (Sandberger). – Wenz, p. 175. 1917 Strobilops (Strobilops) uniplicata (Sandberger). – Wenz, p. 69. 1918 Strobilops uniplicata (Sandberger). – Wenz, p. 20. 1923 Strobilops (Strobilops) uniplicata uniplicata (A. Braun). – Wenz, p. 1057 (cum syn.). 1927 S. [trobilops] uniplicata (Sdbg.). – Pilsbry, p. 47, pl. 8, figs 10–13. 1948 S. [trobilops] uniplicata (Sandberger). – Pilsbry, p. 865, text-figs 468/10–13. 1999 Discostrobilops uniplicatus (Braun). – Esu, p. 331. 1999b Strobilops uniplicata (A. Braun, 1851). – Stworzewicz, p. 156, figs 44, 45. 2009 S.[trobilops] (Discostrobilops) uniplicatus (Braun, 1851). – Moser et al., p. 47. non 1891 Gasterodonta uniplicata Braun sp. – Penecke, p. 358 (= Strobilops planus Clessin, 1885). Material. – 2 specimens (NHMW 2013/0572/0023), 199 specimens (NHMW 1909/0001/0019). Figure 6. A–D – Mastus complanatus (Reuss in Reuss & Meyer, 1849), NHMW 2013/0572/0018. • E – Mastus filocinctus (Reuss, 1861), NHMW 2013/0572/0019. • F – Mastus? filocinctus (Reuss, 1861), NM-PM-P 753. • G – Mastus filocinctus (Reuss, 1861), NM-PM-P 752. • H, I – Paracoryna diezi (Flach, 1891), NHMW 2013/0572/0020. • J – Paracoryna diezi (Flach, 1891) (protoconch of H). • K, L – Strobilops (Eostrobilops) elasmodonta (Reuss, 1861), NHMW 2013/0572/0022. • M – Strobilops (Eostrobilops) elasmodonta (Reuss, 1861) (protoconch of K). • N–Q – Strobilops (Discostrobilops) uniplicatus (Braun in Walchner, 1851), NHMW 2013/0572/0023. • R–V – Strobilops (Eostrobilops) fischeri (Wenz in Fischer & Wenz, 1914), NHMW 2013/0572/0021. • W – Strobilops (Eostrobilops) fischeri (Wenz in Fischer & Wenz, 1914) (protoconch of S). 844 Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake B A C D F E G N R I H T S K O U P V L J M Q W 845 Bulletin of Geosciences • Vol. 89, 4, 2014 Dimensions. – Diameter: 2.2 mm, height: 0.95 mm (Fig. 6U); diameter: 2.25 mm, height: 1.0 mm (Fig. 6V). Discussion. – In contrast to the other Strobilopsidae from the Most Basin, this species has a slightly malleated protoconch. As stated by Klika (1891), many of the Bohemian shells are slightly flatter, have a wider umbilicus and a thinner peristome compared with typical shells from the Mainz Basin. In addition, Wenz (1915a) observed that the infraparietal fold disappears earlier in the shells from Tuchořice. These features, however, are quite variable in our material and do not allow a convincing separation of the Bohemian shells as distinct species. Rupelian shells from Dvérce, tentatively referred to as Strobilops uniplicatus by Klika (1891), represent most probably a different species as indicated already by Klika himself, referring to the much finer and irregular ribs. Occurrence. – A widespread species, which is known in the Most Basin only from Tuchořice. Wenz (1923) lists numerous Late Oligocene and Early Miocene occurrences from Germany (e.g. Hochheim, Wiesbaden, Mainz, Budenheim, Thalfingen, and Donaurieden). This species was also mentioned from the Early Miocene of Bełchatów and Szczerców (Poland) by Stworzewicz (1999b) and Wagner & Matl (2007). 1918 Pyramidula (Gonyodiscus) falcifera (Boettger). – Wenz, p. 12. 1923 Pleurodiscus falcifera (Boettger). – Wenz, p. 1069 (cum syn.). 1930 Goniodiscus falcifera Böttg. – Pfeffer, p. 43. ? 1933 Pleurodiscus falciferus (O. Boettger). – Wenz, p. 8. 1935 Pleurodiscus falciferus (Bttg.). – Pilsbry, p. 179. 1999 Pleurodiscus falciferus (Boettger). – Esu, p. 332. Material. – 2 specimens (NHMW 2013/0572/0024), 17 specimens (NHMW 1909/0001/0023). Dimensions. – Diameter: 13 mm, height: 7.5 mm. Discussion. – Wenz (1923) and Pilsbry (1935) discussed this species to be probably conspecific with Pleurodiscus orbicularis (Klein, 1846). The descriptions and illustrations of that Early Miocene species as given in Klein (1846) and Quenstedt (1884), however, are very poor. Jooss (1918) emended the definition of that species, referring to it as “Pyramidula (Gonyodiscus) orbicularis Klein emend. Jooss”. Following Jooss, that species differs from the Bohemian species clearly in the sculpture of the last whorl, which comprises amalgamated and weak axial ribs persisting on the base. Confusion with the rather smooth Hyalinia orbicularis Klein sensu Sandberger, 1875, which is placed in Aegopinella by Lueger (1981), is impossible. Family Pleurodiscidae Wenz, 1923 Type species. – Helix Balmei Potiez & Michaud, 1838; original designation. Recent, Southern Europe. Occurrence. – Known from Tuchořice and Korozluky. Occurrences from the Early Miocene of Thalfingen, Hauchenberg at Missen, Eggingen and Ulm in Germany, listed by Wenz (1923, 1933), need verification and might partly represent Pleurodiscus orbicularis. Pleurodiscus falciferus (Boettger, 1870) Figure 7A–D Family Valloniidae Morse, 1864 Genus Pleurodiscus Wenz, 1919 *1870a Helix (Patula) falcifera Boettg.; Boettger, p. 288, pl. 13, figs 3a–d. 1891 Patula falcifera Boettger. – Klika, p. 36, text-figs 28a–d. 1892 Patula falcifera Boettger. – Klika, p. 35, text-figs 28a–d. 1911 [Patula] falcifera Bttg. – Kafka, p. 67. 1916 Patula falcifera Boettger. – Thuma, p. 83. 1917 Pyramidula (Gonyodiscus) falcifera (Boettger). – Wenz, p. 55. Genus Vallonia Risso, 1826 Type species. – Vallonia rosalia Risso, 1826 [= Vallonia pulchella (Müller, 1774)]; by monotypy. Recent, France. Vallonia lepida (Reuss in Reuss & Meyer, 1849) Figure 7L–Q 1849a H.[elix] lepida Rss.; Reuss in Reuss & Meyer, p. 12 (nomen nudum). Figure 7. A, B–D – Pleurodiscus falciferus (Boettger, 1870), NHMW 2013/0572/0024. • E, F – Esuinella nana (Braun in Walchner, 1851), NHMW 2013/0572/0027. • G – Esuinella nana (Braun in Walchner, 1851) (protoconch of E). • H–K – Spermodea plicatella (Reuss in Reuss & Meyer, 1849), NHMW 2013/0572/0028. • L–O – Vallonia lepida (Reuss in Reuss & Meyer, 1849), NHMW 2013/0572/0025. • P, Q – Vallonia lepida (Reuss in Reuss & Meyer, 1849) (protoconch of L). • R, S – Acanthinula tuchoricensis (Klika, 1891), NHMW 2013/0572/0026. • T – Acanthinula tuchoricensis (Klika, 1891) (protoconch of S). • U – Acanthinula sp., NM-PM-P 448. • V–X – Granaria intrusa (Slavík, 1869), NHMW 2013/0572/0029. 846 Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake A D B C M E L H N J I F O S U T R G P K Q V W X 847 Bulletin of Geosciences • Vol. 89, 4, 2014 *1849b H.[elix] lepida m.; Reuss in Reuss & Meyer, p. 24, pl. 2, fig. 4. 1861 H.[elix] lepida Reuss. – Reuss, p. 68. 1869a Helix (Glaphyra) lepida Rss. – Slavík, p. 244, pl. 4, figs 1, 2. 1869b Helix (Glaphyra) lepida Rss. – Slavík, p. 261, pl. 4, figs 1, 2. 1875 Helix (Vallonia) lepida. – Sandberger, p. 375, pl. 22, figs 16–16b. 1891 Helix (Vallonia) lepida Reuss. – Klika, p. 44, text-fig. 37. 1892 Helix (Vallonia) lepida Reuss. – Klika, p. 43, text-fig. 37. 1911 [Helix] (Vallonia) lepida Rss. – Kafka, p. 67. 1916 Helix (Vallonia) lepida Reuss. – Thuma, p. 83. 1917 Vallonia lepida (Reuss). – Wenz, p. 70. 1923 Vallonia lepida lepida (Reuss). – Wenz, p. 903. 1996 Vallonia lepida (Reuss, 1849). – Gerber, p. 88, figs 3d, 29a–g, 31a–f, 32a–c, 33. 2008 Vallonia lepida (Reuss, 1849). – Prysjazhnjuk, p. 95, pl. 3, fig. 23, pl. 6, figs 51, 52. 2013 Vallonia lepida (Reuss, 1849). – Salvador, p. 14, figs 15, 16. Material. – 2 specimens (NHMW 2013/0572/0025), 16 specimens (NHMW 1909/0001/0032). Dimensions. – Diameter: 2.6 mm, height: 1.4 mm; diameter: 2.3 mm, height: 1.6 mm. Discussion. – As already discussed by Harzhauser et al. (2014), the enormous stratigraphic and geographic range of this species as given by Gerber (1996) seems unlikely. Morphologically, however, no distinction was possible so far (see Gerber 1996). The protoconch of the Bohemian species displays a prominent sculpture consisting of numerous spiral threads developing from a granulose initial part. This sculpture is much weaker in Middle and Late Miocene specimens, as described by Harzhauser & Kowalke (2002), Harzhauser & Binder (2004) and Harzhauser et al. (2014). This indicates either variability in protoconch sculpture or the presence of another distinct species, such as Vallonia subpulchella (Sandberger, 1875). As Vallonia lepida is based on Early Miocene specimens from Korozluky, the identity of the herein described specimens is without any doubt. Occurrence. – Vallonia lepida (Reuss in Reuss & Meyer, 1849) sensu Gerber (1996) appears during the Oligocene and persists up to the Pliocene. Geographically it was distributed from France to China; the record of Mongolia by Stworzewicz (2007) refers to another species as revised by Neubauer et al. (2013). In the Most Basin it is known from Korozluky, Tuchořice and Lipno. 848 Genus Acanthinula Beck, 1847 Type species. – Helix aculeata Müller, 1774; subsequent designation by Albers & Martens (1860). Recent, Europe. Acanthinula tuchoricensis (Klika, 1891) Figure 7R–T 1870a Patula sp. nov. – Boettger, p. 288. *1891 Helix (Acanthinula) tuchořicensis n.; Klika, p. 42, text-figs 35a–c. 1892 Helix (Acanthinula) tuchořicensis Klika. – Klika, p. 41, text-figs 35a–c. 1911 [Helix (Acanthinula)] Tuchořicensis Kl. – Kafka, p. 67. 1917 Acanthinula tuchoricensis (Klika). – Wenz, p. 69. 1923 Acanthinula tuchoricensis (Klika). – Wenz, p. 978 (cum syn.). 1972 Helix (Acanthinula) tuchoricensis Klika. – Schlickum & Truc, p. 189. non 1902b Acanthinula tuchoricensis Klika. – Andreae, p. 10 [= Pupa (Modicella) trochulus Sandberger, 1875]. non 1911 Theba (Acanthinula) tuchoricensis Klika. – Gaál, p. 66, pl. 2, fig. 5 [= Pupa (Modicella) trochulus Sandberger, 1875]. non 1944 Acanthinula tuchoricensis Klika. – Troll-Obergfell, p. 383 [= Pupa (Modicella) trochulus Sandberger, 1875]. non 2006 Acanthinula tuchoricensis (Klika), 1891. – Kókay, p. 70, pl. 25, fig. 15. Material. – 2 specimens (NHMW 2013/0572/0026), 200 specimens (NHMW 1909/0001/0030). Dimensions. – Diameter: 1.7 mm, height: 2.0 mm (Fig. 7R); diameter: 1.5 mm, height: 1.6 mm (Fig. 7S). Description. – This stout, pupoid-conical species is characterised by its densely spaced prosocline axial threads and wide-spaced ribs. A much weaker spiral sculpture appears between the axial elements, developing a nearly cancellate pattern on the last whorl. Its protoconch has a slightly immersed initial part followed by a bulbous, convex whorl with prominent spiral threads. Discussion. – These features are reminiscent of the Late Miocene Acanthinula trochulus (Sandberger, 1875), which differs in its broad conical shell. Similarly, the Oligocene A. paludinaeformis (Sandberger, 1858) and the Pliocene A. clairi Schlickum & Truc, 1972 are distinguished by their more conical outlines. Occurrence. – Only known from Tuchořice and Lipno. The Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake shells from the Middle Miocene of Austria, Poland and Hungary described as Acanthinula tuchoricensis by Troll-Obergfell (1944), Andreae (1902b) and Gaál (1911) represent A. trochulus (see also Wenz 1923, Stworzewicz 1999b). The Early Miocene whorl fragment from the Hungarian Somlóvásárhely drilling, identified as A. tuchoricensis by Kókay (2006), differs clearly in its straight primary ribs and the absence of spiral sculpture. Acanthinula sp. Figure 7U 1925 Helix (Acanthinula) paludinaeformis Sandb. var. – Petrbok, p. 1, unnumbered pl., fig. 7. Material. – 1 specimen (NM-PM-P 448). Dimensions. – Diameter 2.7 mm, height: 3.5 mm. Discussion. – The only available specimen was described by Petrbok (1925) as unnamed variety of Acanthinula paludinaeformis (Sandberger, 1875). This Oligocene species, as described by Wenz (1914) from the Oligocene of Germany, has a distinctly lower and more conical spire. The specimen differs also from the frequent A. tuchoricensis in its much larger size and higher, more convex last whorl but agrees well in spire outline and macro-sculpture. Therefore, it might represent a new species. As we do not have any information on micro-sculpture and protoconch morphology of the rather poorly preserved specimen, we list this species in open nomenclature. Occurrence. – Only known from Tuchořice. Genus Esuinella Harzhauser, Neubauer & Georgopoulou gen. nov. Type species. – Helix nana Braun in Walchner, 1851 (Oligocene, Germany). Etymology. – In honour of Daniela Esu, palaeontologist at the University of Rome and well-known specialist for Neogene molluscs. Diagnosis. – Minute valloniid consisting of 1.25 bulbous protoconch whorls with strongly malleated sculpture and ca 2.5 convex teleoconch whorls with deeply incised sutures. Sculpture consisting of prosocline axial ribs, separated by broad interspaces with faint axial threads, crossed by weak spiral furrows. The rather high (for confamiliar taxa), conical spire is contrasted by a broad last whorl with a convex adapical part separated by a weak angulation from the weakly convex lower part of the whorl. Aperture semicircular; thin peristome; outer lip is attached to the lower third of the last whorl; deep and moderately wide umbilicus. Discussion. – Helix nana was revised by Falkner (1974), and was placed in Planogyra Morse, 1864, which is based on the extant American species Helix asteriscus Morse, 1857. This decision was based on comparison with shells of the type species but especially on comparisons with shells of the European Vallonia astoma Boettger, 1909 (considered as Planogyra by Falkner 1974). The latter species is now generally considered to be a junior synonym of Helix sororcula Benoit, 1859 (e.g. Welter-Schultes 2012), which is the type species of Gittenbergia Giusti, Castagnolo & Manganelli, 1985. Falkner (1974) recognised the huge stratigraphic gap between “Helix” nana and the extant species and discussed the possibility that a (sub)generic separation might be necessary. His main argument to maintain the fossil species in the Recent genus was the similarity of the malleated protoconch of “Helix” nana and “Vallonia astoma” and the idea to indicate a phylogenetic relation by applying a broad genus concept. Despite this superficial similarity, the protoconch of Gittenbergia sororcula is less bulbous, consists of more whorls (1.5 versus 1.25), which are only moderately convex and its sculpture is weaker. The spire is distinctly lower than in E. nana and the predominating last whorl is regularly convex, resulting in a Vallonia-like outline. The umbilicus is wider and the outer lip is attached to the middle of the last whorl but placed distinctly lower in Esuinella nana. The similar extant SW European Plagyrona palucida (Shuttleworth, 1852) differs in its immersed protoconch and its striate sculpture (Cianfanelli et al. 2012). Thus, aside from similarities in sculpture, the Oligocene and Miocene genus Esuinella has little in common with extant European valloniids and spelaeodiscinids. Included species. – Helix nana Braun in Walchner, 1851 (Oligocene, Germany). Occurrence. – The new genus Esuinella is only known from the Late Oligocene to Middle Miocene of Central Europe. Esuinella nana (Braun in Walchner, 1851) comb. nov. Figure 7E–G *1851 Helix nana A. Braun; Braun in Walchner, p. 1140, p. 56 in offprint, No. 340. 849 Bulletin of Geosciences • Vol. 89, 4, 2014 ? 1869a Helix (Hyalina) euristhmia sp. nov.; Slavík, p. 248, pl. 4, figs 5, 6. ? 1869b Helix (Hyalina) euristhmia sp. nov.; Slavík, p. 263, pl. 4, figs 5, 6. 1875 Patula (Acanthinula) nana A. Braun. – Sandberger, p. 374, pl. 22, figs 14–14c. 1891 Helix (Acanthinula) nana A. Braun. – Klika, p. 41, text-figs 34a–c. 1892 Helix (Acanthinula) nana A. Braun. – Klika, p. 40, text-figs 34a–c. 1902b Acanthinula nana (Al. Braun). – Andreae, p. 10. 1911 Helix (Acanthinula) nana Br. – Kafka, p. 67. 1914 Acanthinula nana (Sdbg.). – Wenz in Fischer & Wenz, p. 195, pl. 7, fig. 32. 1917 Acanthinula nana (Sandberger). – Wenz, p. 69. 1923 Acanthinula nana (A. Braun). – Wenz, p. 972. 1974 Planogyra nana. – Falkner, p. 240, pl. 10, fig. 4, pl. 11, figs 9, 10. 1999b Planogyra nana (A. Braun, 1851). – Stworzewicz, p. 149, figs 32, 33. 2006 Planogyra nana (A. Braun), 1843. – Kókay, p. 71, pl. 26, figs 3, 4. Material. – 2 specimens (NHMW 2013/0572/0027), 841 specimens (NHMW 1909/0001/0029). Dimensions. – Diameter: 1.15 mm, height: 0.85 mm (Fig. 7E); diameter: 1.25 mm, height: 0.9 mm (Fig. 7F). Description. – This tiny species has a very characteristic shape with depressed conical spire and broad last whorl with a slight angulation in its upper third. It has a strongly convex, malleated protoconch of 440 μm diameter. The teleoconch sculpture starts abruptly, consisting of strongly prosocline, sharp and raised axial ribs with 3–5 weaker secondary threads and intercalations of various tertiary axial threads crossed by delicate spiral furrows. The axial elements fade out on the base where the spiral furrows become dominant. Discussion. – The Bohemian specimens are probably conspecific with the Oligocene Helix nana Braun in Walchner, 1851 from Hochheim-Flörsheim in Germany (e.g. Wenz 1923), based on the illustrations and descriptions in Sandberger (1875). Some doubts may arise from the fact that the microscopic spiral sculpture was never mentioned for the Oligocene shells. Unfortunately, we were not able to study any topotypic specimens from Hochheim. Similarly, Falkner (1974) investigated only shells from Tuchořice and Zwiefaltendorf but did not study Oligocene nana-specimens. Wenz (1923) treated Helix (Hyalina) euristhmia Slavík, 1869 as synonym of H. nana. The illustrations of this species in Slavík (1869a, b) are extremely poor and show a rather turbinoid shell. 850 Occurrence. – Esuinella nana is known from the Late Oligocene of Hochheim-Flörsheim in Germany and the Early Miocene of Tuchořice, Bełchatów in Poland (Stworzewicz 1999b) and Somlóvásárhely in Hungary (Kókay 2006), as well as from the Middle Miocene of Opole in Poland and from Undorf and Zwiefaltendorf in Germany (Falkner 1974). Genus Spermodea Westerlund, 1902 Type species. – Helix lamellata Jeffreys, 1830; subsequent designation by Pilsbry (1922–1926). Recent, Great Britain. Spermodea plicatella (Reuss in Reuss & Meyer, 1849) Figure 7H–K 1849a H.[elix] plicatella Rss.; Reuss in Reuss & Meyer, p. 11 (nomen nudum). *1849b H.[elix] plicatella m.; Reuss in Reuss & Meyer, p. 21, pl. 1, fig. 10. 1861 H.[elix] paludinaeformis A. Br. – Reuss, p. 63 (non Helix paludinaeformis Sandberger, 1858). 1861 H.[elix] plicatella Reuss. – Reuss, p. 63. 1891 Helix (Acanthinula) plicatella Reuss. – Klika, p. 43, text-figs 36a–c. 1892 Helix (Acanthinula) plicatella Reuss. – Klika, p. 42, text-figs 36a–c. 1911 [Helix (Acanthinula)] plicatella Rss. – Kafka, p. 67. 1914 Acanthinula plicatella (Rss.). – Wenz in Fischer & Wenz, p. 105, pl. 7, fig. 31. 1916 Acanthinula plicatella (Reuss). – Wenz, p. 174. 1917 Acanthinula plicatella (Reuss). – Wenz, p. 69. 1923 Acanthinula plicatella (Reuss). – Wenz, p. 974 (cum syn.). 1972 Spermodea plicatella. – Schlickum & Truc, p. 189. 1974 [Spermodea] plicatella (Reuss). – Falkner, p. 233 (pars). ? 1977 Spermodea plicatella (Reuss). – Moayedpour, p. 64, pl. 4, figs 8, 9; non figs 2–5. 1981 Spermodea plicatella (Reuss). – Lueger, p. 35. Material. – 2 specimens (NHMW 2013/0572/0028), 1913 specimens (NHMW 1909/0001/0031). Dimensions. – Diameter: 1.85 mm, height: 1.5 mm (Fig. 7I); diameter: 2.0 mm, height: 1.6 mm (Fig. 7H). Description. – The protoconch consists of 1.25 moderately convex whorls and is covered by a dense and irregular groove-ridge pattern, causing a low-relief malleated surface. The sculpture of the teleoconch consists of regularly spaced, thin axial ribs with very faint axial threads in the broad interspaces. Weak and densely spaced spiral Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake grooves are most prominent on the base and the lower part of the last whorl close to the periphery, indenting the axial ribs. Discussion. – This species is among the most common gastropods in the Most Basin. Based on this rich material, it becomes obvious that the shape and especially the axial ribbing display only moderate variability. This is in contrast to the statement of Falkner (1974), who included also Middle Miocene specimens from Opole in this species. These stratigraphically much younger specimens from Poland might represent a separate species. The poorly preserved specimens from the Early Miocene of Theobaldshof/Rhön illustrated by Moayedpour (1977, pl. 4, figs 8, 9) might indeed represent S. plicatella although the depressed spire would be very untypical. The specimen illustrated by Moayedpour (1977, pl. 4, figs 3–6), however, differ considerably from S. plicatella in the somewhat angulated last whorl and the very weak sculpture on the base. Spermodea plicatella is reminiscent of the Pliocene S. demarcqui Schlickum & Truc, 1972, which differs mainly in its higher shell and the delicate protoconch sculpture. A separation based on shell size as discussed by Schlickum & Truc (1972) is not possible. The stratigraphically closer Middle Miocene Spermodea candida Falkner, 1974 differs clearly in its narrower umbilicus and the much denser axial ribbing. Occurrence. – Aside from Tuchořice, Wenz (1923), Falkner (1974) and Moayedpour (1977) cite Spermodea plicatella also from the Oligocene of Hochheim-Flörsheim and Hoppetenzell (Germany), the Early Miocene of Donaurieden, Theobaldshof/Rhön (Germany), and the Middle Miocene of Opole (Poland). As discussed above, most of these occurrences need verification. Family Chondrinidae Steenberg, 1925 Genus Granaria Held, 1837 Type species. – Pupa frumentum Draparnaud, 1801; subsequent designation by Herrmannsen (1847). Recent, Europe. Granaria intrusa (Slavík, 1869) Figure 7V–X 1861 P.[upa] subvariabilis Sandb. – Reuss, p. 70 (non Pupa subvariabilis Sandberger, 1858). *1869a Pupa (Torquilla) intrusa sp. nov.; Slavík, p. 259, pl. 4, figs 12, 13. 1869b Pupa (Torquilla) intrusa sp. nov.; Slavík, p. 267, pl. 4, figs 12, 13. 1870a Pupa (Torquilla Stud.) subvariabilis Sandb. – Boettger, p. 295 (non Pupa subvariabilis Sandberger, 1858). 1891 Torquilla intrusa Slavík. – Klika, p. 87, text-fig. 83. 1892 Torquilla intrusa Slavík. – Klika, p. 84, text-fig. 83. 1889 T.[orquilla] intrusa (Slav.). – Boettger, p. 20. 1911 Torquilla intrusa H. [sic]. – Kafka, p. 68. 1917 Torquilla intrusa (Slavik). – Wenz, p. 65. 1923 Abida intrusa (Slavik). – Wenz, p. 944 (cum syn.). 2013 Granaria intrusa (Slavik, 1869). – Höltke & Rasser, p. 191. Material. – 2 specimens (NHMW 2013/0572/0029), 11 specimens (NHMW 1909/0001/0070). Dimensions. – Diameter: 3.1 mm, height: 8.8 mm (Fig. 7V); diameter: 3.1 mm, height: 8.4 mm (Fig. 7W, X). Description. – This broad-fusiform shell is characterised by its conical spire and the nearly conical last three whorls; the last whorl develops a marked, sometimes even keel-like, angulation on the base. The broad-ovoid aperture bears a very prominent angular tooth, a thickened parietal tooth, two close-set columellar folds and 3–4 palatal folds. The latter appear deep in the aperture and aside from the prominent lower one are very weak; especially the fourth uppermost one is almost completely reduced in some species. Discussion. – This species was recently transferred to Granaria by Höltke & Rasser (2013) in their revision of Miocene Chondrinidae. Boettger (1870a) and Sandberger (1875) considered this species as synonym of Granaria subvariabilis (Sandberger, 1863) from the Oligocene of the Mainz Basin. Later, this idea was rejected by Klika (1891), Wenz in Fischer & Wenz (1914) and even Boettger (1889) himself. Despite the similarities concerning the apertural features, Granaria intrusa can be distinguished from G. subvariabilis based on its larger size, by having more teleoconch whorls and especially due to its characteristic angulation close to the base of the last whorl. Erroneously, Wenz (1923, p. 948) listed “Torquilla subvariabilis Klika” in his synonymy of Granaria subvariabilis, although Klika (1891) clearly described the Bohemian shells as Torquilla intrusa (Klika in turn erroneously listed Pupa subvariabilis Sandberger in his synonymy of T. intrusa, although he discussed the differences between both species in great detail). This confusion might have deduced Höltke & Rasser (2013) to list Granaria subvariabilis also from Tuchořice. Occurrence. – Only known from Tuchořice and Lipno. 851 Bulletin of Geosciences • Vol. 89, 4, 2014 Family Vertiginidae Fitzinger, 1833 of the apertural wall coincides with the position of the lower palatal plica. Genus Vertigo Müller, 1774 Type species. – Vertigo pusilla Müller, 1774; by monotypy. Recent, Europe. Vertigo minor Boettger, 1870 Figure 8A–D *1870a [Vertigo callosa] Varietät minor m.; Boettger, p. 296, pl. 13, fig. 7. 1889 V.[ertigo] (Alaea) minor. – Boettger, p. 305. 1891 Vertigo (Alaea) minor Boettger. – Klika, p. 97, text-fig. 92. 1892 Vertigo (Alaea) minor Boettger. – Klika, p. 93, text-fig. 92. 1911 [Vertigo (Alaea)] minor Bttg. – Kafka, p. 68. 1917 Vertigo (Alaea) minor Boettger. – Wenz, p. 67, unnumbered figure on p. 68. 1919 Vertigo minor Boettger. – Pilsbry & Cooke, p. 216. 1923 Vertigo (Vertigo) minor Boettger. – Wenz, p. 996 (cum syn.). 1999b Nesopupa minor (Boettger, 1870) comb. nov.; Stworzewicz, p. 144, figs 21, 22 (figs 19, 20?). Material. – 3 specimens (NHMW 1909/0001/0078). Dimensions. – Diameter: 1.16 mm, height: 1.5 mm; diameter: 1.05 mm, height: 1.45 mm (Fig. 8A); diameter: 1.05 mm, height: 1.45 mm (Fig. 8C). Description. – A tiny, very stout ovoid shell comprising 1.5 high and convex protoconch whorls with deep sutures and granulose-malleated surface. This sculpture is crossed by several weak spiral furrows which are best developed on the late part of the protoconch and in its adapical half. The teleoconch consists of 2.5 convex whorls with incised sutures; the first 1.5 whorls bear a dense sculpture of prominent, regularly spaced prosocline axial ribs; these become much weaker and irregular on the high last whorl where they are crossed by very indistinct, broad spiral threads producing a faint pattern of shallow pits in the interspaces and low nodes at the intersections. Umbilicus narrow and deep; U-shaped aperture with strong and protruding columellar lamella; very large and bent parietal tooth accompanied by a distinctly smaller and thinner angular tooth and an indistinct infraparietal denticle deep in the aperture; two widely spaced, large but narrow palatal plicae and a well-developed basal denticle. A knob-like low suprapalatalis may occur deep in the aperture. The more prominent lamellae are all characterised by strongly convex, tube-like inflated margins delimitating much thinner inner walls. A shallow sulcus on the outer surface 852 Discussion. – Stworzewicz (1999b) was the first to recognise the unusual shell sculpture of this tiny vertiginid and suggested a placement in Nesopupa Pilsbry, 1900a within the subfamily Nesopupinae Steenberg, 1925. Indeed, Nesopupa sp. of Gargominy (2008, fig. 2A) is strikingly similar to Vertigo minor concerning sculpture and shape of the teleoconch but develops a sheet-like angularis. The protoconch of Nesopupa as described by Gargominy (2008) agrees with the Bohemian shell in its granulose surface but lacks any striae. These, however, are developed on the protoconch of the nesopupine genus Nesoropupa Gargominy, 2008 (type species Nesoropupa duodecim Gargominy, 2008). Nesoropupa differs from Vertigo minor in having a much higher and more convex protoconch, which lacks the malleated surface (see Gargominy 2008, figs 3C, D). Another similar (sub)genus is Nesodagys Cooke & Pilsbry in Pilsbry & Cooke, 1920. Nesodagys thaanumi (Ancey, 1904) develops a similar aperture like the Bohemian species with near-identical parietal and palatal plicae and a comparable columellaris and agrees in the tube-like margins of the lamellae. A close relation to any of these genera, however, seems unlikely as they represent extant Indo-West Pacific groups. Therefore, these similarities are most probably convergences and we follow Manganelli et al. (2008) and Nordsieck (written comm. 2014) and place this species in Vertigo based on its apertural features, although the protoconch sculpture is rather untypical for European Vertigo species. The characteristic sculpture and aperture allow a clear separation from the Oligocene–Early Miocene Pupa trigonostoma Sandberger, 1863 and Vertigo blumi Boettger, 1884. These species develop at least one additional whorl – thus being distinctly larger – and bear prominent growth lines on the entire teleoconch. Since Boettger (1889) and Pilsbry & Cooke (1920, p. 379), both species were “traditionally” placed in Nesopupa (Stworzewicz 1999b, Manganelli et al. 2008). In our opinion, it is questionable if both are even congeneric: differences are the absence of palatal plicae in “Pupa” trigonostoma and the conspicuous continuation of the aperture margin into the angular lamella in “Vertigo” blumi. Representatives of the Oligocene–Miocene genus Ptychalaea Boettger, 1889, which is also considered to be a Nesopupinae by Pilsbry & Cooke (1920), differ clearly in their smooth shells. Stworzewicz (1999b) considered the shells from the Middle Miocene of Bełchatów to be conspecific with those from the Early Miocene of the Most Basin. As already noticed by Stworzewicz (1999b), the Polish specimens differ from those from Tuchořice in the shorter angular tooth and especially in the very indistinct basal denticle. In respect to Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake the large stratigraphic gap between both occurrences we consider the Polish specimens to represent a distinct chrono-subspecies. Occurrence. – The species is known from the Burdigalian of Tuchořice. The Middle Miocene (MN 5) specimens from Bełchatów in Poland are probably a younger chronosubspecies. Vertigo angulifera Boettger, 1884 *1884 Vertigo (Alaea) angulifera sp. nov.; Boettger, p. 271, pl. 4, figs 10a–c. 1923 Vertigo (Vertilla) angulifera angulifera Boettger. – Wenz, p. 1006 (cum syn.). 1967 Vertigo (Vertilla) angulifera angulifera O. Boettger. – Schütt, p. 207, fig. 9. 1985 Vertigo (Vertilla) angulifera angulifera Boettger. – Čejchan, p. 176, pl. 1, figs 1–5, pl. 2, figs 1–3. 1998 Vertigo angulifera. – Finger, p. 44, pl. 9, fig. H. 2008 Vertigo angulifera Boettger, 1884. – Harzhauser et al., p. 50, figs 5.7, 5.8. Material. – No material was available; the specimen described by Čejchan (1985) is stored in the collection of the Charles University in Prague. Dimensions. – Diameter: 0.85 mm, height: 1.45 mm. Discussion. – Vertigo angulifera is a very rare species in the Most Basin; the only record was documented by Čejchan (1985). Occurrence. – The distribution of this species was summarised by Stworzewicz (1999b) and Harzhauser et al. (2014): it appears during the Early Miocene (Bełchatów, Frankfurt, Tuchořice, Dolnice), is recorded from the Early/Middle Miocene of Undorf (Germany) and the Middle Miocene Badenian of the Rein Basin (Austria) and Hungary and persists up the late Middle Miocene Sarmatian (Gratkorn Basin and North-Alpine Foreland Basin in Austria, Steinheim Basin in Germany). During the Middle Miocene it spreads as far east as the Fore-Caucasus. Vertigo callosa (Reuss in Reuss & Meyer, 1849) Figure 8F–H 1849a Vertigo callosa Rss.; Reuss in Reuss & Meyer, p. 11 (nomen nudum). *1849b V.[ertigo] callosa m.; Reuss in Reuss & Meyer, p. 30, pl. 3, fig. 7. 1861 P.[upa] callosa Reuss. – Reuss, p. 72. 1891 Vertigo (Alaea) callosa Reuss. – Klika, p. 96, text-figs 91a, b. 1892 Vertigo (Alaea) callosa Reuss. – Klika, p. 91, text-figs 91a, b. 1875 Pupa (Vertigo) callosa Reuss. – Sandberger, p. 400, pl. 24, figs 19–19b. 1889 Vertigo (Alaea) callosa (Rss.). – Boettger, p. 71, pl. 7, figs 3, 4. 1916 Vertigo (Alaea) callosa Reuss. – Thuma, p. 84. 1916 Vertigo (Alaea) callosa (Reuss). – Wenz, p. 174. 1917 Vertigo (Alaea) callosa (Reuss). – Wenz, p. 67. 1923 Vertigo (Vertigo) callosa (Reuss). – Wenz, p. 983 (cum syn.). 1964 Vertigo callosa (Reuss, 1849). – Čtyroký et al., table for p. 149. 1977 Vertigo (Vertigo) callosa (Reuss). – Moayedpour, p. 63, pl. 4, figs 1, 2. 1999b Vertigo callosa (Reuss, 1849). – Stworzewicz, p. 137, figs 6, 7. 2008 Vertigo callosa (Reuss, 1849). – Prysjazhnjuk, p. 92, pl. 3, figs 20, 21, pl. 5, figs 37–43. 2013 Vertigo callosa (Reuss, 1849). – Salvador, p. 15, figs 17, 18. Material. – 2 specimens (NHMW 2013/0572/0030); >8,600 specimens (NHMW 1909/0001/0077, 0079). Dimensions. – Diameter: 1.15 mm, height: 1.75 mm (Fig. 8H); diameter: 1.22 mm, height: 1.85 mm (Fig. 8G). Description. – Small ovoid shells with cyrtoconoid spire, large last whorl and domical, moderately convex protoconch lacking any sculpture. The three teleoconch whorls are moderately convex and bear prosocline growth lines. Two aperture types are represented in the rich material. The dominant type bears 6 teeth with two prominent parietal teeth, a large columellar fold, a knob-like basal denticle and two palatal teeth with a stronger lower one (Fig. 8F, H). Rarely a small infraparietal denticle may appear. The second, less abundant group comprises shells with an additional weak suprapalatal denticle and a very low, usually twofold basal knob (Fig. 8G). A comparable intra-specific variability was described by Stworzewicz (1999b) from Bełchatów in Poland, who rejects a taxonomic separation of both morphotypes. Occurrence. – A widespread species during the Early and Middle Miocene in central and eastern Europe, ranging to the Caucasus region; latest occurrences are known from the Late Miocene of Austria and Hungary (see Stworzewicz 1999b and Harzhauser et al. 2014 for details). In the Most Basin it is known from Korozluky, Tuchořice, Lipno and the Kralupy drillings. 853 Bulletin of Geosciences • Vol. 89, 4, 2014 Vertigo tuchoricensis Pilsbry in Pilsbry & Cooke, 1919 Figure 8I–J 1861 P.[upa] microstoma Reuss; Reuss, p. 73, pl. 2, fig. 8 (non Pupa microstoma Pfeiffer, 1854). 1869a Vertigo microstoma Reuss. – Slavík, p. 260 (non Pupa microstoma Pfeiffer, 1854). 1869b Vertigo microstoma Reuss. – Slavík, p. 272 (non Pupa microstoma Pfeiffer, 1854). 1875 Pupa (Vertigo) microstoma Reuss. – Sandberger, p. 438, pl. 24, figs 17–17b (non Pupa microstoma Pfeiffer, 1854). 1891 Vertigo microstoma Reuss. – Klika, p. 98, text-figs 93a, b (non Pupa microstoma Pfeiffer, 1854). 1892 Vertigo microstoma Reuss. – Klika, p. 93, text-figs 93a, b (non Pupa microstoma Pfeiffer, 1854). 1911 [Vertigo (Alaea)] microstoma Rss. – Kafka, p. 68 (non Pupa microstoma Pfeiffer, 1854). *1919 Vertigo tuchoricensis new name; Pilsbry in Pilsbry & Cooke, p. 218. 1923 Vertigo (Vertigo) tuchoricensis Pilsbry. – Wenz, p. 1002 (cum syn.). Material. – One specimen (NHMW 2013/0572/0031). Reuss (1861) do not match at all. He describes two prominent palatal plicae of which the upper one is stronger and in the middle of the aperture, whilst the weaker, lower one is close to the columella. Further, Reuss (1861) mentioned two weaker but acute parietal lamellae. The illustration shows an additional, small columellar fold and an enormous angularis but no palatal ones. Reuss (1861) emphasised the narrow, half-moon shaped aperture with thin peristome and based the (preoccupied) name microstoma on this feature. Concerning the illustration, we are inclined to ascribe this unusual aperture to the fact that parts of the peristome may simply be broken off. By introducing tuchoricensis as replacement name, Pilsbry in Pilsbry & Cooke (1919) settled the nomenclatorial problem, but no author was able to present new data on the morphology. Our specimen agrees with the original description largely in its internal apertural features and the strong convexity of the spire whorls. Thus, although uncertainties remain, we tentatively consider it as Vertigo tuchoricensis. This species is clearly separated from other vertiginids of the Most Basin by its smaller size, the strongly convex whorls, the granulose protoconch, the narrow umbilicus and the wide aperture with rather fragile and thin elements. Dimensions. – Diameter: 1.0 mm, height: 1.6 mm. Occurrence. – Only known from Lipno and Tuchořice, Description. – Very small and thin-shelled species; the high-domical protoconch is weakly granulated; this microsculpture is best developed on the adapical part and becomes much weaker towards the lower suture. The teleoconch sculpture consists of weakly prosocline, somewhat irregular and broad growth lines. The spire whorls are strongly convex, separated by deep sutures; the umbilicus is narrow and partly covered by the inner lip. Peristome only moderately thickened and wide with two prominent but very narrow parietal folds and an also narrow columellar fold. No basal denticle is visible in our specimen; the lower palatal tooth is covered by sediment; above follow a comparatively prominent upper palatal tooth and a distinct but small suprapalatal denticle. Discussion. – The identity of this species is completely unclear. It is based on a lost specimen illustrated and described by Reuss (1861), who did not provide any measurements. Aside from the problem that the name Pupa microstoma was preoccupied, the description and illustration of Genus Negulus Boettger, 1889 Type species. – Pupa reinhardti Jickeli, 1874; original designation. Recent, Ethiopia. Negulus suturalis (Sandberger, 1858) Figure 8K–N *1858 Pupa suturalis A. Braun; Sandberger, p. 54, pl. 6, figs 2–2a. 1861 P.[upa] suturalis A. Br. sp. – Reuss, p. 71. 1875 Pupa lineolata A. Braun. – Sandberger, p. 397 (pars). 1891 Negulus lineolatus Al. Braun. – Klika, p. 89, text-figs 85a, b (non Pupa lineolata A. Braun, 1875). 1892 Negulus lineolatus Al. Braun. – Klika, p. 85, text-figs 85a, b (non Pupa lineolata A. Braun, 1875). 1889 Negulus lineolatus (Al. Br.). – Boettger, p. 45 (pars). Figure 8. A–C – Vertigo minor Boettger, 1870, NHMW 1909/0001/0078. • D – Vertigo minor Boettger, 1870, protoconch of A, B. • E – Ptychalaea flexidens (Reuss, 1861), NHMW 1909/0001/0076. • F – Vertigo cf. callosa (Reuss in Reuss & Meyer, 1849), NHMW 2013/0572/0030. • G, H – Vertigo callosa (Reuss in Reuss & Meyer, 1849) NHMW 2013/0572/0030. • I, J – Vertigo tuchoricensis Pilsbry in Pilsbry & Cooke, 1919, NHMW 2013/0572/0031. • K–M – Negulus suturalis (Sandberger, 1858), NHMW 2013/0572/0032. • N – Negulus suturalis (Sandberger, 1858) (protoconch of L). • O–Q – Negulus raricosta (Slavík, 1869), NHMW 2013/0572/0033. 854 Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake A F G E C B H I K L D P O M J N Q 855 Bulletin of Geosciences • Vol. 89, 4, 2014 1911 Negulus lineolatus A. B. – Kafka, p. 69 (non Pupa lineolata A. Braun, 1875). 1914 Negulus suturalis (Sandberger). – Wenz, p. 92, pl. 5, fig. 13. 1916 Negulus suturalis (Sandberger). – Wenz, p. 172. 1917 Negulus suturalis (Sandberger). – Wenz, p. 65. 1923 Negulus suturalis suturalis (Sandberger). – Wenz, p. 1024. 1993 Negulus suturalis (Sandberger, 1858). – Stworzewicz, p. 398. 1999b Negulus suturalis (Sandberger, 1858). – Stworzewicz, p. 146, figs 23–25. 2002 Negulus suturalis (Sandberger, 1858). – Binder, p. 168, pl. 1, fig. 10, text-figs 6a, b. 2006 Negulus suturalis suturalis (Sandberger), 1858. – Kókay, p. 61, pl. 21, figs 12, 13. Material. – 2 specimens (NHMW 2013/0572/0032), 840 specimens (NHMW 1909/0001/0071). Dimensions. – Diameter: 0.75 mm, height: 1.5 mm (Fig. 8L); diameter: 0.65 mm, height: 1.45 mm (Fig. 8K). Discussion. – A very common species at Tuchořice. Stworzewicz (1999b) presented a detailed description of this species with emphasis on the microsculpture and its shell variability. The Bohemian shells agree largely with those from Bełchatów aside from the slightly weaker peristome. Occurrence. – A wide-spread species known from the Oligocene of Hochheim-Flörsheim (Germany), the Early Miocene of Donaurieden, Erbach, Budenheim (Germany), Bełchatów (Poland), Teiritzberg (Austria), Somlóvásárhely (Hungary) and the Middle Miocene of Opole and Bełchatów (Poland). In the Most Basin it was detected at Tuchořice (Wenz 1923; Stworzewicz 1993, 1999b; Binder 2002; Kókay 2006). ? 1993 Negulus raricostatus [sic] (Slavik, 1869). – Stworzewicz, p. 399. 1999 Negulus raricostatus [sic] (Slavik). – Esu, p. 332. Material. – 2 specimens (NHMW 2013/0572/0033), 5 specimens (NHMW 1909/0001/0072). Dimensions. – Diameter: 1.5 mm, height: 2.25 mm (Fig. 8O); diameter: 1.55 mm, height: 2.25 mm (Fig. 8P). Description. – The shell bears wide and regularly spaced, strongly oblique prosocline and sharp axial ribs with faint growth lines in the interspaces. A dense pattern of spiral grooves appears in the interspaces, crossing the growth lines without intersecting the axial ribs. The convex protoconch has a slightly immersed initial part and develops a coarse, malleated granulose sculpture. The apertural margin is thin; only the inner lip is slightly expanded. Discussion. – This species was originally described as Pupa raricosta by Slavík (1869) and erroneously listed as raricostatus by Wenz (1917, 1923); the correct name is Negulus raricosta. It is much rarer than the congeneric Negulus suturalis from which it differs very distinctly in its stout shape with strongly convex whorls. Occurrence. – Known only from Tuchořice; the much younger occurrences from the Middle Miocene of Opole in Poland mentioned by Andreae (1902b) need confirmation. Subfamily Truncatellininae Steenberg, 1925 Genus Truncatellina Lowe, 1852 Type species. – Pupa linearis Lowe, 1852; by monotypy. Recent, Madeira. Negulus raricosta (Slavík, 1869) Figure 8O–Q *1869a Pupa (Pupilla) raricosta sp. nov.; Slavík, p. 258, pl. 4, figs 9–11. 1869b Pupa (Pupilla) raricosta sp. nov.; Slavík, p. 266, pl. 4, figs 9–11. 1891 Negulus raricosta Slavík. – Klika, p. 91, text-figs 86a–c. 1892 Negulus raricosta Slavík. – Klika, p. 86, text-figs 86a–c. ? 1902b Negulus raricostatus [sic] Slav. – Andreae, p. 17. 1911 [Negulus] raricosta Sl. – Kafka, p. 69. 1917 Negulus raricostatus [sic] (Slavik). – Wenz, p. 66. 1923 Negulus raricostatus [sic] (Slavik). – Wenz, p. 1024 (cum syn.). 856 Truncatellina splendidula (Sandberger, 1875) Figure 9E–H 1849a Pupa minutissima Hartm. – Reuss in Reuss & Meyer, p. 11 (non Pupa minutissima Hartmann, 1821). 1849b P.[upa] minutissima Hartm. – Reuss in Reuss & Meyer, p. 29, pl. 3, fig. 6 (non Pupa minutissima Hartmann, 1821). 1861 P.[upa] cryptodus A. Br.? – Reuss, p. 71 (non Pupa cryptodus Sandberger, 1858). 1869a Pupa (Pupilla) cryptodus A. Braun. – Slavík, p. 267 (non Pupa cryptodus Sandberger, 1858). *1875 P.[upa] splendidula Sandb.; Sandberger, p. 397. 1891 Isthmia splendidula Sandberger. – Klika, p. 93, text-figs 88a, b. Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake 1892 Isthmia splendidula Sandberger. – Klika, p. 89, text-figs 88a, b. 1889 Isthmia splendidula (Sbgr.). – Boettger, p. 48 in offprint, pl. 6, fig. 9. 1911 Isthmia splendidula Sndb. – Kafka, p. 69. 1914 Isthmia splendidula (Sdbg.). – Wenz in Fischer & Wenz, p. 93, pl. 5, fig. 15. 1917 Isthmia splendidula (Sandberger). – Wenz, p. 66. 1923 Truncatellina splendidula (Sandberger). – Wenz, p. 1021 (cum syn.). Material. – 2 specimens (NHMW 2013/0572/0035), 266 specimens (NHMW 1909/0001/0074). Dimensions. – Diameter: 0.75 mm, height: 1.55 mm (Fig. 9F); diameter: 0.7 mm, height: 1.45 mm (Fig. 9G–H). Description. – High-cylindrical shell with smooth protoconch, moderately convex, nearly smooth whorls, incised sutures, and narrow umbilicus. Aperture U-shaped, with a prominent columellar fold, a slightly weaker parietal lamella and a knob-like palatal denticle deep in the aperture. In contrast to the ovoid aperture in the schematic illustration of Boettger (1889), the outer lip is nearly straight (see also Fischer & Wenz 1914). Discussion. – Wenz (1923) mentioned also Truncatellina cryptodus (Sandberger, 1858) from Tuchořice with question mark. He referred to Slavík (1869a), who did not mention this species anymore in the descriptive part of Slavík (1869b). Occurrence. – This species was described from the Late Oligocene of Hochheim-Flörsheim (Germany) and the Early Miocene of Budenheim and Frankfurt am Main (Germany). In the Most Basin it is only known from Tuchořice. Family Gastrocoptidae Pilsbry, 1918 Genus Gastrocopta Wollaston, 1878 Subgenus Albinula Sterki, 1892 Type species. – Pupa contracta Say, 1822; subsequent designation by Pilsbry (1916). Recent, North America. Gastrocopta (Albinula) turgida (Reuss in Reuss & Meyer, 1849) Figure 9A–D 1849a Vertigo turgida Rss.; Reuss in Reuss & Meyer, p. 11 (nomen nudum). *1849b V.[ertigo] turgida m.; Reuss in Reuss & Meyer, p. 30, pl. 3, fig. 8. 1861 P.[upa] turgida Reuss. – Reuss, p. 71. 1889 L.[eucochilus] quadriplicatum (Al. Br.) var. lamellidens Sbgr. – Boettger, p. 56. 1891 [Leucochilus quadriplicatum A. Braun] var. lamellidens Sandberger. – Klika, p. 92, text-figs 87a, b. 1892 [Leucochilus quadriplicatum A. Braun] var. lamellidens Sandberger. – Klika, p. 88, text-figs 87a, b (non Pupa lamellidens Sandberger, 1858). 1911 Leucochilus quadriplicatum A. Br. var. lamellidens Sndb. – Kafka, p. 69. 1916 Leucochilus quadriplicatum A. Braun, var. lamellidens Sandberger. – Thuma, p. 84. 1916 Leucochila turgida (Reuss). – Wenz, p. 173. 1916/17 Vertigo turgida Reuss. – Pilsbry, p. 115. 1917 Leucochila turgida (Reuss). – Wenz, p. 66. 1918 Leucochila turgida (Reuss). – Wenz, p. 20. 1923 Gastrocopta (Albinula) turgida turgida (Reuss). – Wenz, p. 922 (cum syn.). 1999 Gastrocopta (Albinula) turgida (Reuss). – Esu, p. 331. 1999b Gastrocopta turgida (Reuss, 1849). – Stworzewicz, p. 159, figs 51–54. 2000a Vertigo turgida Reuss, 1849. – Manganelli & Giusti, p. 73. 2000a Gastrocopta (Albinula) quadriplicata (Braun, 1851). – Manganelli & Giusti, pl. 8, figs 1, 2. ? 2008 Gastrocopta (Albinula) turgida (Reuss, 1849). – Prysjazhnjuk, p. 89, pl. 3, figs 15–17, pl. 4, figs 26–28. Material. – 2 specimens (NHMW 2013/0572/0034), 104 specimens (NHMW 1909/0001/0104, 1909/0001/0104). Dimensions. – Diameter: 1.7 mm, height: 2.6 mm (Fig. 9A–C); diameter: 1.7 mm, height: 2.6 mm (Fig. 9D). Description. – This species is characterised by its sheetlike anguloparietal lamella with undulating margin, joining the apertural margin. The aperture bears three palatal plicae with a well-developed lower one and a usually still recognisable upper palatal plica, whereas the suprapalatal denticle is strongly reduced. The columellar lamella is emerging, semicircular and obliquely ascending. The entire aperture and especially the anguloparietal lamella are covered by tiny granulae. Prosocline growth lines are distinct but never as strong as indicated by the illustration of Reuss in Reuss & Meyer (1849). Discussion. – This species is very common, easily recognisable and well represented in the NHMW collection. Therefore, we have little doubts that our shells represent the species described by Reuss in Reuss & Meyer (1849) 857 Bulletin of Geosciences • Vol. 89, 4, 2014 although the original illustration is very poor. The poor illustration and vague description of Reuss in Reuss & Meyer (1849) was base for much confusion on its relation to Gastrocopta (Albinula) quadriplicata (Braun in Walchner, 1851) and Gastrocopta (Albinula) lamellidens (Sandberger, 1858). Manganelli & Giusti (2000a) identified specimens from Tuchořice as Gastrocopta (Albinula) quadriplicata and considered G. turgida as potential senior synonym of G. lamellidens Sandberger, 1858. In our opinion, because only a single but frequent Gastrocopta species is represented in the material from the Most Basin, Gastrocopta (Albinula) quadriplicata sensu Manganelli & Giusti (2000a) might rather be a junior synonym of P. turgida. Occurrence. – Known from Korozluky, Tuchořice and Lipno in the Most Basin. Further Early Miocene occurrences are listed by Wenz (1923) from France (Noaillan, Mérignac, Saucats) and Germany (Hochheim-Flörsheim, Donaurieden, Öpfingen, Ehingen, Lehr bei Ulm, Eggingen). Stworzewicz (1999b) documented G. turgida also from the Early and early Middle Miocene of Bełchatów (Poland). The species is probably also recorded from several Oligocene localities but the unclear status of Gastrocopta lamellidens (Sandberger, 1858) and G. quadriplicata (Braun in Walchner, 1851) does not allow a clear statement. The occurrence of this species in the Early Miocene of Sjenica in Trijebine (Serbia) is doubtful as it is based only on internal molds (Prysjazhnjuk 2008). A Middle Miocene record from Opole in Poland listed by Wenz (1923) was identified as G. edlaueri (Wenz, 1921) by Stworzewicz (1999b). 1919 V.[ertigo] flexidens Reuss. – Pilsbry, p. 157. 1923 Vertigo (Ptychalaea) flexidens flexidens (Reuss). – Wenz, p. 1003 (cum syn.). non 1891 Pupa (Vertigo) flexidens Rss. – Penecke, p. 368 (= Leucochilus quadriplicatum mut. suevica Boettger, 1889). Material. – One specimen (NHMW 1909/0001/0076). Dimensions. – Diameter: 1.2 mm, height: 1.75 mm. Discussion. – The identity of this species has been unlear due to the very poor descriptions and drawings of Reuss (1861) and Sandberger (1875), who both present a rather conical shell with rapidly contracting base and heartshaped aperture. A part of the peristome might have been broken off, resulting in the strange aperture shape. In fact, the aperture of the stout ovoid shell is more or less U-shaped with wide peristome as already indicated by Klika (1891) and Boettger (1889). The characteristic feature is the prominent and slightly bent angularis, which extends to the apertural margin. The likewise strong and bent parietal tooth is adjoined by a weak infraparietal denticle. Similarly, the two prominent palatal teeth are adjoined by a pointed suprapalatal denticle, which causes a trefoil-like sulcus in the aperture together with the angular tooth. The basal denticle is weak and knob-like; the columellar fold is slightly weaker than the palatal and parietal teeth. According to G. Manganelli (written comm. 2014) Ptychalaea should be placed in the Gastrocoptidae based on the morphology of the angularis, which is tentatively followed herein. Genus Ptychalaea Boettger, 1889 Type species. – Pupa flexidens Reuss, 1861; original designation. Early Miocene, Czech Republic. Ptychalaea flexidens (Reuss, 1861) Figure 8E *1861 P.[upa] flexidens Reuss; Reuss, p. 74, pl. 2, fig. 9. 1875 Pupa (Vertigo) flexidens Reuss. – Sandberger, p. 439, pl. 24, figs 18–18b. 1889 Vertigo (Ptychalaea) flexidens (Rss.). – Boettger, p. 69, pl. 7, fig. 1. 1891 Vertigo (Ptychalaea Boettg.) flexidens Reuss. – Klika, p. 95, text-figs 90a, b. 1892 Vertigo (Ptychalaea Boettg.) flexidens Reuss. – Klika, p. 90, text-figs 90a, b. 1911 Vertigo (Ptychalaea) flexidens Rss. – Kafka, p. 69. 1916 Vertigo (Ptychalaea) flexidens (Reuss). – Wenz, p. 173. 1917 Vertigo (Ptychalaea) flexidens (Reuss). – Wenz, p. 67. 858 Occurrence. – This species is known from Tuchořice and Lipno. Wenz (1923) mentions several additional occurrences from the Early Miocene of Germany (e.g. Öpfingen, Budenheim, Wiesbaden, Donaurieden). Occurrences of this species in the Middle Miocene Rein Basin in Austria as mentioned by Penecke (1891) actually represent Gastrocopta suevica (Boettger 1889, Harzhauser et al. 2014). Family Orculidae Pilsbry, 1918 Genus Nordsieckula Harl & Harzhauser gen. nov. Type species. – Pupa subconica Sandberger, 1858; Late Oligocene, Germany. Etymology. – In honour of Hartmut Nordsieck (Senckenberg Museum, Frankfurt), the leading specialist for European Tertiary terrestrial gastropods. Diagnosis. – Medium-sized orculid of barrel-like shape Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake with narrow umbilicus; 7–9 weakly convex whorls; sculpture of protoconch smooth, teleoconch weakly or moderately ribbed; apex rather rounded, not attenuate. First whorls regularly increasing in diameter, last whorls of similar width. Aperture one-sided rounded, the outer margin curved stronger, margins not closing. Aperture armouring: parietalis strong; subangularis bulge-like and weakly expressed; two equally strong upper columellar lamellae extend to the aperture margins; a third, bulge-like columellar lamella (subcolumellaris) sometimes present at aperture margin, but not entering the shell; palatal plica missing. Discussion. – Pupa subconica was traditionally placed in Orcula Held, 1837, with the extant type species Pupa dolium Draparnaud, 1801. The genus Orcula was revised by Harl et al. (2011) and Páll-Gergely et al. (2013), who emphasised the difficulties to distinguish the various extant orculid genera and subgenera based on conchological data alone. Nevertheless, the characteristic bulge-like subangularis (sensu Hausdorf 1996) separates N. subconica from extant species of Orcula, as already discussed by Sandberger (1858) and Boettger (1889). N. subconica is probably closest related to the Middle Miocene N. falkneri (Hausdorf, 1995), which is known from Middle Miocene sediments of Gündlkofen (Germany) and Nowa Wies Królewska (Poland) (Hausdorf 1995). Nordsieckula falkneri and N. subconica strongly resemble each other in shell form and aperture characteristics, including the shared presence of a subangularis. The similarities between these two species let us suggest that N. subconica (or related taxa) is ancestral to O. falkneri, wherefore the latter species is also placed in Nordsieckula herein. The third columellar lamella (subcolumellaris) is a distinctive trait shown by N. subconica only, but not by N. falkneri or any of the extant Orcula species. Generally, the shell characteristics of N. subconica and N. falkneri indicate a close relationship with the orculid genera Orcula, Orculella, Sphyradium and Schileykula. Some of the Orculella species from Northern Africa (Libya) also exhibit a subangularis (Brandt 1956). Included species. – Pupa subconica Sandberger, 1858 and Orcula falkneri Hausdorf, 1995 are placed in Nordsieckula herein. Occurrence. – Nordsieckula is known from the Late Oligocene to the Middle Miocene of Central Europe. Nordsieckula subconica (Sandberger, 1858) comb. nov. Figure 9I *1858 Pupa subconica Sandb.; Sandberger, p. 51, pl. 5, figs 7–7c. 1868 Pupa subconica Sandb. – Reuss, p. 82, pl. 1, fig. 3. 1875 Pupa (Orcula) subconica Sandberger. – Sandberger, p. 394, pl. 23, figs 8–8c. 1889 Orcula subconica (Sbgr.). – Boettger, p. 14. 1891 Orcula subconica Sandberger. – Klika, p. 88, text-figs 84a–c. 1892 Orcula subconica Sandberger. – Klika, p. 84, text-figs 84a–c. 1911 Orcula subconica Sandb. – Kafka, p. 68. 1917 Orcula cf. subconica (Sandberger). – Wenz, p. 64. 1922 Orcula subconica (Sandberger). – Pilsbry, p. 3, pl. 4, fig. 10. 1923 Orcula subconica (Sandberger). – Wenz, p. 1033 (cum syn.). Material. – One fragmentary specimen (NHMW 1868/0014/0002) from Tuchořice and one complete shell from Hochheim in Germany (NHMW 1865/0011/0049) (Fig. 9I). A single fragment stored in the Prague collection (NM-PM-P 1094) consists only of few spire whorls and is unidentifiable. Much more material is stored in the Senckenberg Museum in Frankfurt (Tuchořice: SMF 151718/1; Hochheim: SMF 151713/1, SMF 151714, SMF 151715/2, SMF 151716/2, SMF 151717/3, SMF 185029). Dimensions. – Hochheim specimen: diameter: 3.6 mm, height: 6.2 mm. Discussion. – This species seems to be very rare at Tuchořice. Reuss (1868) mentioned four poorly preserved specimens with largely filled apertures. Boettger (1889) confirmed the identification based on a single specimen in his collection. Later, Wenz (1917), based on the same specimen, doubted that the Bohemian shell is fully conspecific with the Oligocene species due to the stronger axial sculpture. The specimen illustrated by Reuss (1868) is still present in the NHMW collection. It consists of 7 spire whorls, which agree in every detail with the specimens from the type locality Hochheim. A difference in axial sculpture, as proposed by Wenz (1917), is definitely not present. Investigations of the rich collection of the Senckenberg Museum by J.H. confirmed that the specimens from both occurrences are conspecific. Occurrence. – This species is known from the Late Oligocene of Hochheim-Flörsheim (Germany) and from the Burdigalian of Tuchořice. Clade Elasmognatha Mörch, 1864 Superfamily Succineoidea Beck, 1837 Family Succineidae Beck, 1837 Subfamily Succineinae Beck, 1837 859 Bulletin of Geosciences • Vol. 89, 4, 2014 Genus Oxyloma Westerlund, 1885 Type species. – Succinea hungarica Hazay, 1880 [= Oxyloma dunkeri (Pfeiffer, 1865)], by monotypy. Recent, Hungary. Oxyloma affinis (Reuss in Reuss & Meyer, 1849) comb. nov. Figure 9K–N 1849a S.[uccinea] affinis Rss.; Reuss in Reuss & Meyer, p. 11 (nomen nudum). 1849a Succinea Pfeifferi Rossm. – Reuss in Reuss & Meyer, p. 11 (non Succinea Pfeifferi Rossmässler, 1835). 1849b S.[uccinea] Pfeifferi Rossm. – Reuss in Reuss & Meyer, p. 18, pl. 1, fig. 2 (non Succinea Pfeifferi Rossmässler, 1835). *1849b S.[uccinea] affinis m.; Reuss in Reuss & Meyer, p. 18, pl. 1, fig. 3. 1855 S.[uccinea] Pfeifferi Rossm. – Pictet, p. 31, pl. 57, fig. 26 (non Succinea Pfeifferi Rossmässler, 1835). 1861 S.[uccinea] affinis Reuss. – Reuss, p. 63. 1861 S.[uccinea] Pfeifferi Rossm.? – Reuss, p. 63 (non Succinea Pfeifferi Rossmässler, 1835). 1875 Succinea (Amphibina) affinis Reuss. – Sandberger, p. 440, pl. 24, figs 21–21b. 1875 Succinea peregrina Sandberger; Sandberger, p. 440, pl. 24, figs 22–22b. 1891 Succinea peregrina Sandberger. – Klika, p. 99, text-figs 94a, b. 1891 Succinea affinis Reuss. – Klika, p. 99, text-figs 95a, b. 1892 Succinea peregrina Sandberger. – Klika, p. 94, text-figs 94a, b. 1892 Succinea affinis Reuss. – Klika, p. 95, text-figs 95a, b. 1911 Succinea peregrina Sndbg. – Kafka, p. 68. 1911 Succinea affinis Rss. – Kafka, p. 68. 1916 Succinea peregrina Sandberger. – Thuma, p. 84. 1917 Succinea (Amphibina) affinis Reuss. – Wenz, p. 72. 1917 Succinea (Amphibina) peregrina Sandberger. – Wenz, p. 72. 1923 Succinea (Amphibina) affinis Reuss. – Wenz, p. 890 (cum syn.). 1923 Succinea (Amphibina) peregrina peregrina Sandberger. – Wenz, p. 895 (cum syn.). specimens (NHMW 1847/0032/0087, 1909/0001/0081, 1909/0001/0080); 4 specimens in the Prague collection (NM-PM-P 586-587 = Succinea peregrina sensu Klika, 1891, NM-PM-P 578 = Succinea affinis sensu Klika, 1891). Dimensions. – Diameter: 5.0 mm, height: 9.0 mm (Fig. 9K, L); diameter: 4.8 mm, height: 8.5 mm (Fig. 9M, N). Discussion. – Since Reuss in Reuss & Meyer (1849a), the Succineidae from Tuchořice have traditionally been split into two species. A stouter morphotype with low spire, moderately convex whorls, more or less straight columella, wide basal aperture, prominent growth lines and faint spiral grooves was originally identified with the extant Succinea pfeifferi Rossmässler, 1835 [= Oxyloma elegans (Risso, 1826)]. For this morphotype, Sandberger (1875) introduced Succinea peregrina as new name. Shells with higher spire, more convex whorls, slightly concave columella, narrower aperture and overall more delicate sculpture were described as Succinea affinis Reuss in Reuss & Meyer, 1849. Most authors, such as Sandberger (1875) and Klika (1891), seem to have reproduced the original description of Reuss uncritically. Only Wenz (1917) discussed the possibility that both taxa are morphotypes of a single species. Moreover, both taxa have been recorded so far exclusively from the Most Basin. Within the herein studied material, which comprises more than 120 shells of all ontogenetic stages, none of the above mentioned features qualified as useful criterion. All combinations of the supposed discriminating features appear in the population. Interestingly, many of the fully grown shells show injuries that occurred during early growth stages. Subsequent shell growth was clearly influenced by these injuries, often resulting in more pronounced spiral sculpture and/or a wider last whorl. Therefore, we treat both taxa as a single species for which Succinea affinis is the first available name. Occurrence. – Only known from Korozluky, Tuchořice and Lipno. Informal group Sigmurethra Pilsbry, 1900b Superfamily Clausilioidea Gray, 1855 Note. – The generic assignments of species within this superfamily follow Nordsieck (1981a, b, 2000, 2013a). Material. – 5 specimens (NHMW 2013/0572/0036), 123 Figure 9. A–D – Gastrocopta turgida (Reuss in Reuss & Meyer, 1849), NHMW 2013/0572/0034. • E – Truncatellina splendidula (Sandberger, 1875), NHMW 2013/0572/0035 (protoconch of H). • F–H – Truncatellina splendidula (Sandberger, 1875), NHMW 2013/0572/0035. • I – Nordsieckula subconica (Sandberger, 1858), NHMW 1865/0011/0049 (note that this specimen is from Hochheim, Germany). • J – Pseudocalaxis? insignis (Babor, 1897), NM-PM-P 450. • K–N – Oxyloma affinis (Reuss in Reuss & Meyer, 1849), NHMW 2013/0572/0036. • O, P – Palaeoglandina gracilis (von Zieten, 1832), NHMW 2013/0572/0040. • Q–T – Pseudoleacina oligostropha (Reuss in Reuss & Meyer, 1849), NHMW 2013/0572/0041. • U–X – Pseudoleacina producta (Reuss in Reuss & Meyer, 1849), NHMW 2013/0572/0042. 860 Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake A D B E F C H G L K Q R I N M S T U J O P V W X 861 Bulletin of Geosciences • Vol. 89, 4, 2014 Family Filholiidae Wenz, 1923 Genus Triptychia Sandberger, 1875 Type species. – Clausilia antiqua von Zieten, 1832; subsequent designation by Wenz (1923). Early Miocene, Germany. Triptychia vulgata (Reuss in Reuss & Meyer, 1849) Figure 10A–D 1849a Clausilia vulgata Rss.; Reuss in Reuss & Meyer, p. 11 (nomen nudum). *1849b C.[lausilia] vulgata m.; Reuss in Reuss & Meyer, p. 34, pl. 4, fig. 1. 1861 Cl.[ausilia] vulgata Reuss. – Reuss, p. 74, pl. 2, fig. 10. 1875 Clausilia vulgata Reuss. – Sandberger, p. 434, pl. 24, figs 13–13b. 1877 Clausilia (Triptychia) vulgata Reuss. – Boettger, p. 23. 1891 Triptychia (Plioptychia) vulgata Reuss. – Klika, p. 76, text-figs 73a, b. 1892 Triptychia (Plioptychia) vulgata Reuss. – Klika, p. 73, text-figs 73a, b. 1911 Triptychia (Plioptychia) vulgata Rss. – Kafka, p. 68. 1914 Clausilia (Triptychia) vulgata Reuss. – Frankenberger, p. 156. 1914 [Clausilia (Triptychia) vulgata Reuss] f. grandis m. – Frankenberger, p. 157. 1916 Triptychia (Plioptychia) vulgata Reuss. – Thuma, p. 84. 1917 Triptychia [sic] (Plioptychia) vulgata. – Wenz, p. 61. 1923 Triptychia (Plioptychia) vulgata (Reuss). – Wenz, p. 801 (cum syn.). 2006 Triptychia (Plioptychia) vulgata (Reuss 1849). – Schnabel, p. 175, pl. 10, figs 116–119 (cum syn.). non 1967 Triptychia (Plioptychia) vulgata (Reuss). – Schütt, p. 215 [= Triptychia (Plioptychia) schuetti Schnabel, 2006]. Material. – 2 specimens (NHMW 2013/0572/0037), 11 specimens (NHMW 1890/0013/0411, 1909/0001/0063). Dimensions. – Diameter: 4.3 mm, height: 18.0 mm (Fig. 10A, B); diameter: 4.5 mm, height: 17.5 mm (Fig. 10C, D). Description. – This species was discussed in great detail by Schnabel (2006). It is characterised by its moderately stout fusiform shape, a very regular sculpture of densely spaced and sharp axial ribs, a narrow sinulus, a very prominent and protruding parietal lamella (almost perpendicular to the shell axis) and two less prominent, oblique columellar lamellae. 862 Discussion. – This species is the type species of Plioptychia Boettger, 1877, which is based on the palatal plicae of the penultimate whorl, which are developed as transverse folds (Schnabel 2006). As pointed out by Nordsieck (2013a) transitions from normal to transverse folds may occur and therefore the subgeneric status of Plioptychia is not accepted. The Middle Miocene (Sarmatian) shells from Hollabrunn in Austria, identified as Triptychia (Plioptychia) vulgata by Schütt (1967), represent a different species (Schnabel 2006). Occurrence. – Only known from the Most Basin, where it is documented from Korozluky, Tuchořice, Lipno and Pyšná. Family Clausiliidae Gray, 1855 Genus Serrulella Nordsieck in Zilch, 1978 Type species. – Serrulina truci Nordsieck, 1972; original designation. Pliocene, Germany. Serrulella schwageri (Boettger, 1877) Figure 10E–H 1861 Cl.[ausilia] denticulata Reuss; Reuss, p. 76, pl. 2, fig. 12, pl. 3, fig. 12 (non Bulimus denticulatus Olivier, 1801). *1877 Clausilia (Serrulina) Schwageri; Boettger, p. 73. 1891 Serrulina schwageri Boettger. – Klika, p. 79, text-figs 75a, b. 1892 Serrulina schwageri Boettger. – Klika, p. 75, text-figs 75a, b. 1911 [Serrulina] Schwageri Bttg. – Kafka, p. 68. 1914 Clausilia (Serrulina) Schwageri Bttg. – Frankenberger, p. 160. 1917 Serrulina schwageri Boettger. – Wenz, p. 62. 1923 Serrulina schwageri (Boettger). – Wenz, p. 780 (cum syn.). 1972 [Serrulina] schwageri (O. Boettger, 1877). – Nordsieck, p. 172. 1981a [Serrulella] schwageri O. Boettger. – Nordsieck, p. 67. 1981b Serrulella schwageri (O. Boettger). – Nordsieck, p. 107. 1999 Serrulella schwageri (Boettger). – Esu, p. 332. 2000 [Serrulella] schwageri (O. Boettger, 1877). – Nordsieck, p. 2. 2007 [Serrulella] schwageri (O. Boettger, 1877). – Nordsieck, p. 126. Material. – One specimen (NHMW 1909/0001/0065), one specimen (NHMW 1979/2083/0025). Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake Dimensions. – Diameter: 2.0 mm, height: 9.8 mm (Fig. 10E, F); diameter: 2.5 mm, height: 10.5 mm (Fig. 10G, H). Description. – Fusiform shells with weakly convex whorls and moderately detached aperture. Early whorls are smooth; axial ribs appear between the 4th to 6th whorls, becoming successively stronger. The position of the sharp parietal lamella coincides with a deep incision of the adapical part of the aperture. Five about equally spaced parietal and columellar lamellae follow, whilst basal and palatal areas are smooth. Discussion. – Reuss (1861) introduced this species as Clausilia denticulata, overlooking that this name was already preoccupied by the extant Clausilia denticulata (Olivier, 1801). Therefore, Boettger (1877) proposed Clausilia (Serrulina) Schwageri as replacement name. The lack of basal and palatal denticles separates this species from superficially similar clausiliids of the Most Basin such as Serrulastra (Serrulastra) amphiodon (Reuss, 1861) and Serrulastra (Serrustigma) polyodon (Reuss, 1861). Nordsieck (1981a) considers Serrulella andreaei Nordsieck, 1981a from Opole as closely related Middle Miocene species. Occurrence. – Only known from Tuchořice. Genus Serrulastra Nordsieck, 1981 Subgenus Serrulastra Nordsieck, 1981 1981b Serrulastra amphiodon (Reuss). – Nordsieck, p. 107. 1999 Serrulastra amphiodon (Reuss). – Esu, p. 332. 2000 [Serrulastra (Serrulastra)] amphiodon (Reuss, 1860). – Nordsieck, p. 2. 2007 [Serrulastra (Serrulastra)] amphiodon (Reuss, 1860). – Nordsieck, p. 127, pl. 19, fig. 2. Material. – One specimen (NHMW 2013/0572/0038), 20 specimens (NHMW 1909/0001/0066, 1979/2083/0024). Dimensions. – Diameter: 1.75 mm, height: 8.2 mm (Fig. 10I, J); diameter: 1.8 mm, height: 9.0 (Fig. 10K, L). Description. – Somewhat irregular shells with narrow and smooth spire whorls. Only the two adult whorls develop a narrow axial ribbing, which grades into widerspaced, sharp and rarely bifurcate ribs on the cervix. Narrow, elongate aperture with numerous lamellae and denticles of which the parietalis and columellaris are slightly more prominent; only the palatal wall below the sinulus lacks denticles (see Klika 1891 for a detailed description). Discussion. – The small size, the ribbing of the last three teleoconch whorls, the narrow aperture, and the high number of denticles distinguish this species clearly from other clausiliids of the Most Basin. Occurrence. – Only known from Tuchořice and Lipno. Type species. – Clausilia amphiodon Reuss, 1861, original designation. Early Miocene, Czech Republic. Subgenus Serrustigma Nordsieck, 1981 Serrulastra (Serrulastra) amphiodon (Reuss, 1861) Figure 10I–L *1861 Cl.[ausilia] amphiodon Reuss; Reuss, p. 77, pl. 3, fig. 14. 1875 Clausilia (Laminifera) amphiodon Reuss. – Sandberger, p. 436, pl. 24, figs 15–15b. 1877 Clausilia (Serrulina) amphiodon Reuss. – Boettger, p. 72. 1891 Serrulina amphiodon Reuss. – Klika, p. 80, text-figs 76a–c. 1892 Serrulina amphiodon Reuss. – Klika, p. 76, text-figs 76a–c. 1911 [Serrulina] amphiodon Rss. – Kafka, p. 68. 1914 Clausilia (Serrulina) amphiodon Reuss. – Frankenberger, p. 160. 1917 Serrulina amphiodon (Reuss). – Wenz, p. 62. 1923 Serrulina amphiodon (Reuss). – Wenz, p. 777 (cum syn.). 1981a [Serrulastra (Serrulastra)] amphiodon Reuss. – Nordsieck, p. 69. Type species. – Clausilia polyodon Reuss, 1861; original designation. Early Miocene, Czech Republic. Serrulastra (Serrustigma) polyodon (Reuss, 1861) Figure 10M–P *1861 Cl.[ausilia] polyodon Reuss; Reuss, p. 76, pl. 3, fig. 13. 1875 Clausilia polyodon Reuss. – Sandberger, p. 437, pl. 24, figs 16–16c. 1877 Clausilia (Serrulina) polyodon Reuss. – Boettger, p. 293, pl. 13, fig. 5. 1891 Serrulina polyodon Reuss. – Klika, p. 77, text-figs 74a, b. 1892 Serrulina polyodon Reuss. – Klika, p. 74, text-figs 74a, b. 1911 Serrulina polyodon Rss. – Kafka, p. 68. 1914 Clausilia (Serruline [sic]) polyodon Bttg. – Frankenberger, p. 159. 1917 Serrulina polyodon (Reuss). – Wenz, p. 61. 863 Bulletin of Geosciences • Vol. 89, 4, 2014 1923 Serrulina polyodon (Reuss). – Wenz, p. 779 (cum syn.). 1981b Serrulastra (Serrustigma) polyodon (Reuss, 1860). – Nordsieck, p. 98. 1999 S.[errulastra] polyodon (Reuss). – Esu, p. 332. 2000 [Serrulastra (Serrustigma)] polyodon (Reuss, 1860). – Nordsieck, p. 2. 2007 [Serrulastra (Serrustigma)] polyodon (Reuss, 1860). – Nordsieck, p. 127. Material. – One specimen (NHMW 2013/0572/0039), 4 specimens (NHMW 1890/0013/0412, 1909/0001/0064). Dimensions. – Diameter: 2.5 mm, height: 11.7 mm (Fig. 10M, N); diameter: 2.7 mm, height: 11.0 mm (Fig. 10O, P). Description. – Stout fusiform shell characterised by nearly completely smooth surface aside from sharp axial ribs on the cervix. Wide aperture with angulated sinulus and deep incision at the position of the parietal lamella. Parietal and columellar walls bear numerous prominent denticles and lamellae, of which two columellar lamellae are the strongest. Discussion. – This is the type species of Serrustigma Nordsieck, 1981b, which was defined mainly based on its clausilium plate morphology (Nordsieck 1981b). Occurrence. – Only known from Tuchořice and Lipno. Subgenus Serruplica Nordsieck, 1981 Type species. – Clausilia ptycholarynx Boettger, 1877; original designation. Middle Miocene, Czech Republic. Serrulastra (Serruplica) tuchoricensis nom. nov. 1914 Clausilia (Serrulina) ptycholarynx Bttg. var. laevigata m. – Frankenberger, p. 159 (non Clausilia laevigata Potiez & Michaud, 1838). 2007 [Serrulastra (Serruplica)] laevigata Frankenberger. – Nordsieck, p. 127 (non Clausilia laevigata Potiez & Michaud, 1838). Material. – No material was available. Dimensions. – No measurements were provided by Frankenberger (1914); the specimens seem to be comparable to Serrulastra (Serruplica) ptycholarynx (Boettger, 1877), which has an aperture of about 3 mm in height. Discussion. – According to Frankenberger (1914), the species from Tuchořice differs from the Middle Miocene Serrulastra (Serruplica) ptycholarynx (Boettger, 1877) only in the absence of sculpture. As pointed out by Nordsieck (2000, 2007), the name given by Frankenberger (1914) has to be replaced by a new name as it is preoccupied by Clausilia laevigata Potiez & Michaud, 1838. Therefore, we propose Serrulastra (Serruplica) tuchoricensis as new name, referring to the locality Tuchořice. Occurrence. – Only known from Tuchořice. Genus Laminifera Boettger, 1863 Type species. – Clausilia rhombostoma Boettger, 1863; subsequent designation by Wenz (1923) [note that Boettger (1877) did not designate a type species although this is frequently cited in the literature (e.g. Wenz & Zilch 1960)]. Oligocene, Germany. Laminifera mira (Slavík, 1869) Figure 10Q–S *1869a Clausilia (Laminifera) mira sp. nov.; Slavík, p. 255, text-fig. 57. 1869b Clausilia (Laminifera) mira sp. nov.; Slavík, p. 264, text-fig. 57. 1877 Clausilia (Laminifera) mira Slavík. – Boettger, p. 103, pl. 4, figs 41a–c. 1891 Clausilia (Laminifera Boettg.) mira Slavík. – Klika, p. 86, text-figs 82a–c. 1892 Clausilia (Laminifera Boettg.) mira Slavík. – Klika, p. 82, text-figs 82a–c. 1911 [Clausilia] (Laminifera) mira Sl. – Kafka, p. 68. 1914 Clausilia (Laminifera) mira Slavík. – Frankenberger, p. 161. Figure 10. A–D – Triptychia vulgata (Reuss in Reuss & Meyer, 1849), NHMW 2013/0572/0037. • E, F – Serrulella schwageri (Boettger, 1877), NHMW 1909/0001/0065. • G, H – Serrulella schwageri (Boettger, 1877), NHMW 1979/2083/0025. • I, J – Serrulastra (Serrulastra) amphiodon (Reuss, 1861), NHMW 2013/0572/0038. • K, L – Serrulastra (Serrulastra) amphiodon (Reuss, 1861), NHMW 1979/2083/0024. • M, N – Serrulastra (Serrustigma) polyodon (Reuss, 1861), NHMW 2013/0572/0039. • O, P – Serrulastra (Serrustigma) polyodon (Reuss, 1861), NHMW 1909/0001/0064. • Q, R – Laminifera mira (Slavík, 1869), NHMW 1909/0001/0069. • S – Laminifera mira (Slavík, 1869), NHMW 1979/2083/0027. • T, U – Canalicia attracta (Boettger, 1870), NHMW 1909/0001/0068). • V, W – Constricta tenuisculpta (Reuss, 1861), NHMW 1909/0001/0067. • X – Constricta tenuisculpta (Reuss, 1861), NHMW 1979/2083/0028. 864 Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake A C B I J Q R D E L K S M T F O N U G V H P W X 865 Bulletin of Geosciences • Vol. 89, 4, 2014 1917 Laminifera (Laminifera) mira (Slavik). – Wenz, p. 63. 1923 Laminifera (Laminifera) mira (Slavik). – Wenz, p. 798. 1981b Laminifera mira (Slavik). – Nordsieck, p. 107. 1999 Laminifera mira (Slavik). – Esu, p. 332. 2000 [Laminifera] mira (Slavik, 1869). – Nordsieck, p. 3. 2007 [Laminifera (Laminifera)] mira (Slavik, 1869). – Nordsieck, p. 128. non 1915 Clausilia (Laminifera) mira Slavik. – Fischer & Wenz, p. 54, pl. 2, figs 9a, b. non 1977 Laminifera (L.) mira. – Moayedpour, pp. 54, 67. Material. – One specimen (NHMW 1909/0001/0069), 4 specimens (NHMW 1979/2083/0027). Dimensions. – Diameter: 2.0 mm, height: 9.2 mm (Fig. 10Q, R); diameter: 1.8 mm, height: 8.8 mm (Fig. 10S). 2000 “Laminifera” slaviki (Babor, 1897). – Nordsieck, p. 3. 2007 [Baboria] slaviki (Babor, 1897). – Nordsieck, p. 128. Material. – No specimens were available. The type is listed in the digital inventory of the National Museum in Prague but could not be found. Dimensions (after Babor 1897). – Diameter: 2.5 mm, height: 10 mm. Discussion. – Babor (1897) introduced the subgenus Cossmannia for this species, which was preoccupied by Newton (1891). Therefore, Cossmann (1898) proposed Baboria as replacement name. The status of this poorly known genus was accepted recently by Nordsieck (2007). Occurrence. – Only known from Tuchořice. Discussion. – This species is easily recognised by its slender fusiform shape and the widely detached, subcircular to roughly triangular aperture with wide and regularly concave sinulus, broadened margins and narrow, keeled cervix. Two or three weak lamellae appear between the narrow and prominent parietal lamella and the weak columellaris. Occurrence. – Known only from Tuchořice. Fischer & Wenz (1915) and Moayedpour (1977) describe this species also from the Early Miocene of Theobaldshof/Rhön (Germany). The illustration in Fischer & Wenz (1915) shows a species with more slender apical whorls, narrower aperture margins and a stronger parietal lamella. Therefore, these specimens are not conspecific with the Bohemian species (see also Nordsieck 2000). Genus Baboria Cossmann, 1898 Type species. – Clausilia Slavíki Babor, 1897; original designation. Early Miocene, Czech Republic. Baboria slaviki (Babor, 1897) *1897 Clausilia (Cossmannia subsct. n.) Slavíki sp. nov.; Babor, p. 10, text-fig. 4. 1898 Clausilia slaviki. – Cossmann, p. 56. 1914 Clausilia (Laminifera, Baboria) Slaviki Babor. – Frankenberger, p. 162. 1917 Laminifera (Baboria) slaviki (Babor). – Wenz, p. 64. 1923 Laminifera (Baboria) slaviki (Babor). – Wenz, p. 795 (cum syn.). 1981b ? slaviki (Babor, 1897). – Nordsieck, p. 107. 866 Genus Constricta Boettger, 1877 Type species. – Clausilia (Constricta) kochi Boettger, 1877; subsequent designation by Wenz (1923). Oligocene, Germany. Constricta tenuisculpta (Reuss, 1861) Figure 10V–X *1861 Cl.[ausilia] tenuisculpta Reuss; Reuss, p. 75, pl. 2, fig. 11. 1875 Clausilia tenuisculpta Reuss. – Sandberger, p. 435, pl. 24, figs 14–14b. 1877 Clausilia (Constricta) tenuisculpta Reuss. – Boettger, p. 43. 1891 Clausilia (Constricta Bttg.) tenuisculpta Reuss. – Klika, p. 81, text-figs 78a–c. 1892 Clausilia (Constricta Bttg.) tenuisculpta Reuss. – Klika, p. 78, text-figs 78a–c. 1911 [Clausilia (Constricta)] tenuisculpta Rss. – Kafka, p. 68. 1914 Clausilia (Constricta) tenuisculpta Reuss. – Frankenberger, p. 158, text-fig. 2. 1914 Clausilia (Constricta) collarifera Bttg. – Frankenberger, p. 157, text-fig. 1 (non Clausilia collarifera Boettger, 1877). 1917 Constricta tenuisculpta (Reuss). – Wenz, p. 62. 1923 Constricta tenuisculpta (Reuss). – Wenz, p. 770 (cum syn.). 1981b [Constricta] tenuisculpta (Reuss, 1860). – Nordsieck, p. 99. 1993 Constricta tenuisculpta (Reuss, 1861). – Stworzewicz, p. 398. Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake 1999 Constricta tenuisculpta (Reuss). – Esu, p. 332. 2000 [Constricta] tenuisculpta (Reuss, 1860). – Nordsieck, p. 4. 2007 [Constricta] tenuisculpta (Reuss, 1860). – Nordsieck, p. 129, pl. 18, fig. 2. Material. – One specimen (NHMW 1909/0001/0067), 10 specimens (NHMW 1979/2083/0028). Dimensions. – Diameter: 3.5 mm, height: 16.5 mm (Fig. 10V, W); diameter: 3.6 mm, height: 16.9 mm (Fig. 10X). Discussion. – A very characteristic species which is readily distinguished from all other clausiliids of the Most Basin by it large parietal lamella, the strongly protruding columellaris, the wide and smooth apertural margins and the dense and delicate axial ribbing. Occurrence. – Known from Tuchořice; an additional occurrence was mentioned by Stworzewicz (1993) from the Burdigalian of Bełchatów (Poland). A B Figure 11. Cochlodina (Miophaedusa) perforata (Boettger, 1877), holotype, SMF 341913. 2000 Cochlodina (Miophaedusa) perforata (O. Boettger, 1877). – Nordsieck, p. 5. 2007 [Cochlodina (Miophaedusa)] perforata (O. Boettger, 1877). – Nordsieck, p. 130. Material. – One specimen (holotype: SMF 341913). Genus Cochlodina Férussac, 1821 Subgenus Miophaedusa Nordsieck, 1972 Type species. – Clausilia (Dilataria) perforata Boettger, 1877; original designation. Early Miocene, Tuchořice, Czech Republic. Cochlodina (Miophaedusa) perforata (Boettger, 1877) Figure 11A, B Dimensions. – Height: 3.46 mm, width: 2.5 mm. Discussion. – No material seems to have been found of this small species since the description of Boettger (1877). The holotype – the only available apertural fragment – in the collection of the Senckenberg Museum was restudied by Nordsieck (1981a), who confirmed the relation to Cochlodina. Occurrence. – Only known from Tuchořice. *1877 Clausilia (Dilataria) perforata; Boettger, p. 53, pl. 2, figs 25a–f. 1891 Clausilia (Dilataria v. Möllend.) perforata Boettger. – Klika, p. 83, text-figs 79a–f. 1892 Clausilia (Dilataria v. Möllend.) perforata Boettger. – Klika, p. 79, text-figs 79a–f. 1911 [Clausilia] (Dilataria) perforata Bttg. – Kafka, p. 68. 1914 Clausilia (Dilataria) perforata Bttg. – Frankenberger, p. 159. 1917 Dilataria perforata (Boettger). – Wenz, p. 63. 1923 Charpentieria perforata (Boettger). – Wenz, p. 759 (cum syn.). 1972 Miophaedusa perforata. – Nordsieck, p. 166. 1981a Miophaedusa perforata O. Boettger. – Nordsieck, p. 71. 1981b Cochlodina perforata (O. Boettger). – Nordsieck, p. 107. 1999 Cochlodina perforata (Boettger). – Esu, p. 332. Genus Canalicia Boettger, 1863 Type species. – Clausilia articulata (Sandberger, 1863); subsequent designation by Wenz (1923). Oligocene, Germany. Canalicia attracta (Boettger, 1870) Figure 10T, U *1870a Clausilia (Canalicia Boettg.) attracta Boettg.; Boettger, p. 294, pl. 13, figs 5a–d. 1877 Clausilia (Canalicia) attracta Boettger. – Boettger, p. 83. 1891 Clausilia (Canalicia Boettg.) attracta Boettger. – Klika, p. 84, text-figs 81a–d. 1892 Clausilia (Canalicia Boettg.) attracta Boettger. – Klika, p. 81, text-figs 81a–d. 867 Bulletin of Geosciences • Vol. 89, 4, 2014 1911 [Clausilia] (Canalicia?) attracta Bttg. – Kafka, p. 68. 1914 Clausilia (Canalicia) attracta Bttg. – Frankenberger, p. 160. 1916 Clausilia (Canalicia) attracta Boettger. – Thuma, p. 84. 1917 Canalicia attracta (Boettger). – Wenz, p. 63. 1923 Canalicia attracta (Boettger). – Wenz, p. 784. 1981b Canalicia attracta (O. Boettger). – Nordsieck, p. 107. 1999 Canalicia attracta (Boettger). – Esu, p. 332. 2000 [Canalicia] attracta (O. Boettger, 1877). – Nordsieck, p. 5. ? 2006 Canalicia cf. attracta (Boettger), 1870. – Kókay, p. 83, text-fig. 8. 2007 [Canalicia] attracta (O. Boettger, 1877). – Nordsieck, p. 130. Superfamily Achatinoidea Swainson, 1840 Family Ferussaciidae Bourguignat, 1883 Genus Pseudocalaxis Pallary, 1912 Type species. – Pseudocalaxis terebellum Pallary, 1912; subsequent designation by Zilch in Wenz & Zilch (1959). Recent, Egypt. Pseudocalaxis? insignis (Babor, 1897) Figure 9J *1897 Férussacia insignis sp. nov.; Babor, p. 8, text-fig. 3. 1908a Ferussacia insignis Babor. – Pilsbry, p. 218. 1917 Ferussacia (Pseudazeca) insignis Babor. – Wenz, p. 71. 1923 Ferussacia (Ferussacia) insignis Babor. – Wenz, p. 1084 (cum syn.). Material. – One specimen (NHMW 1909/0001/0068). Material. – One specimen (NM-PM-P 450). Dimensions. – Diameter: 2.1 mm, height: 10.9 mm. Dimensions. – Diameter: 4.5 mm, height: 13 mm. Discussion. – Differs from all other clausiliids of the Most Basin in its coarse and partly bifurcate axial ribs, the cord-like basal keel and the canaliculate basal margin. Occurrence. – Known from Korozluky, Tuchořice and Lipno. An aperture fragment from the Lower Miocene Somlóvásárhely drilling in Hungary described by Kókay (2006) might represent an additional occurrence. Canalicia klikai (Babor, 1897) *1897 Clausilia (Serrulina Mousson) Klikai sp. nov.; Babor, p. 14, text-fig. 5. 1914 Clausilia (Canalicia) Klikai Babor. – Frankenberger, p. 161. 1917 Canalicia klikai (Babor). – Wenz, p. 63. 1923 Canalicia klikai (Babor). – Wenz, p. 786. 2000 [Canalicia] klikai (Babor, 1897). – Nordsieck, p. 5. 2007 [Canalicia] klikai (Babor, 1897). – Nordsieck, p. 130. Material. – No material was available. Dimensions (after Babor 1897). – Diameter: 1.5 mm, height: 13 mm. Discussion. – Differs from Canalicia attracta (Boettger, 1870) in its wider aperture with convex left margin. Occurrence. – Only known from Tuchořice. 868 Description. – A single specimen is known so far. It is slender with weakly convex whorls, narrow sutures and glossy surface despite distinct growth lines. The last whorl attains slightly more than half of the total height. Moderately wide, drop-shaped aperture with deeply concave columella, truncated at the base, and a delicate inner lip. The inner parts of the aperture are completely sealed by sediment and glue. Discussion. – Pilsbry (1908a) considered Ferussacia insignis to be related with the extant F. folliculum (Schröter, 1784). This generic affiliation, which was first proposed by Babor (1897), critically discussed by Wenz (1917) and accepted again by Wenz (1923), is very unlikely. The species has little in common with extant species of Ferussacia Risso, 1826, such as F. folliculum and F. carnea (Risso, 1826), which are clearly more bullet-shaped and have a much higher last whorl. Even the very slender Ferussacia lanzarotensis (Mousson, 1872) lacks the deep concavity of the columella. Aside from the morphologic differences, the huge stratigraphic gap between the Early Miocene and the Pliocene, when the first true Ferussacia species appear, raises doubts on the identification. The overall shape of the Bohemian shell is strongly reminiscent of Cecilioides, but differs from species of this genus in its much larger size and the distinctly less bulbous protoconch. Among the extant Ferussaciidae only the African genus Pseudocalaxis agrees insize, shape and apertural features with the Bohemian shell. This generic placement is provisionally and remains problematic, because we cannot exclude that the similarities are convergent. The disjunct geo- Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake graphic occurrence – Miocene in Europe versus Recent in Africa – is no a-priori argument against the identification, as documented by the occurrence of Negulus (van Bruggen, 1994). In any case, it would be unwise to establish a new genus for this species, due to the lack of information on apertural features. Occurrence. – Only known from Tuchořice. Superfamily Testacelloidea Gray, 1840 Family Oleacinidae H. & A. Adams, 1855 shows a very high-spired, elongate shell with acute spire. Similarly, the better drawn specimen from Ulm in Klein (1846) shows a high-spired species. Therefore, it seemed unlikely that the stout Early Miocene Achatina inflata Reuss, 1849 should represent a synonym. Comparisons of specimens from the Mainz Basin and those from the Most Basin by Pfeffer (1930), however, confirmed that both taxa are conspecific as already proposed by Wenz in Fischer & Wenz (1914) and Wenz (1923). Genus Palaeoglandina Wenz in Fischer & Wenz, 1914 Occurrence. – A widespread species in the Late Oligocene and Early Miocene of Germany, Switzerland and France (Wenz 1923); in the Most Basin it is documented from Tuchořice and Lipno. Type species. – Limnaea gracilis von Zieten, 1832; original designation. Early Miocene, Germany. Genus Pseudoleacina Wenz in Fischer & Wenz, 1914 Palaeoglandina gracilis (von Zieten, 1832) Figure 9O–P Type species. – Achatina sandbergeri Thomä, 1845; original designation. Late Oligocene, Germany. *1832 Limnaea gracilis nobis; von Zieten, p. 39, pl. 30, figs 3a, b. 1849a A.[chatina] inflata Rss.; Reuss in Reuss & Meyer, p. 11 (nomen nudum). 1849b A.[chatina] inflata m.; Reuss in Reuss & Meyer, p. 33, pl. 3, fig. 14. 1855 Achatina inflata Reuss. – Pictet, p. 27, pl. 57, fig. 17. 1861 Gl.[andina] inflata Reuss. – Reuss, p. 69. 1875 Glandina inflata Reuss. – Sandberger, p. 408, pl. 21, figs 18, 18a. 1891 Glandina inflata Reuss. – Klika, p. 20, text-fig. 12. 1891 Glandina inflata Reuss. – Maillard, p. 4, pl. 1, fig. 3. 1892 Glandina inflata Reuss. – Klika, p. 20, text-fig. 12. 1911 Glandina inflata Reuss. – Jooss, p. 52. 1911 Glandina inflata Rss. – Kafka, p. 67. 1914 Poiretia (Palaeoglandina) gracilis (Zieten). – Wenz in Fischer & Wenz, p. 38. 1917 Poiretia (Palaeoglandina) gracilis (Zieten). – Wenz, p. 52. 1923 Poiretia (Palaeoglandina) gracilis gracilis (Zieten). – Wenz, p. 839. 1930 Palaeoglandina gracilis (Zieten). – Pfeffer, p. 210, pl. 3, figs 6, 7. 2004 Palaeoglandina inflata (Reuss). – Binder, p. 201. Material. – 2 specimens (NHMW 2013/0572/0040), 9 specimens (NHMW 1896/0034, 1909/0001/0008, 1909/0001/0007). Dimensions. – Diameter: 22 mm, height: 42 mm. Discussion. – The first illustration of Palaeoglandina gracilis from the Early Miocene of Ulm in von Zieten (1832) Pseudoleacina oligostropha (Reuss in Reuss & Meyer, 1849) Figure 9Q–T 1849a A.[chatina] Sandbergeri Thom. – Reuss in Reuss & Meyer, p. 11 (non Achatina Sandbergeri Thomä, 1845). 1849a Achatina oligostropha Rss.; Reuss in Reuss & Meyer, p. 11 (nomen nudum). 1849b A.[chatina] Sandbergeri Thom. – Reuss in Reuss & Meyer, p. 32, pl. 3, fig. 11 (non Achatina Sandbergeri Thomä, 1845). *1849b A.[chatina] oligostropha m.; Reuss in Reuss & Meyer, p. 33, pl. 3, fig. 13. 1861 Gl.[andina] Sandbergeri Thom. – Reuss, p. 70 (non Achatina Sandbergeri Thomä, 1845). 1861 Gl.[andina] oligostropha Reuss. – Reuss, p. 70. 1891 Oleacina neglecta n.; Klika, p. 21, text-fig. 13. 1891 [Oleacina neglecta] var. subcylindrata Bttgr.; Boettger, p. 229. 1892 Oleacina neglecta Klika. – Klika, p. 21, text-fig. 13. 1911 Oleacina neglecta Kl. – Kafka, p. 67. 1914 Poiretia (Pseudoleacina) sandbergeri (Tho.). – Wenz, p. 52 (pars). 1916 Oleacina neglecta Klika. – Thuma, p. 83. 1917 Poiretia (Pseudoleacina) neglecta Klika. – Wenz, p. 52. 1923 Poiretia (Pseudoleacina) oligostropha oligostropha (Reuss). – Wenz, p. 860 (cum syn.). 1923 Poiretia (Pseudoleacina) oligostropha subcylindrata (Boettger). – Wenz, p. 861. 1930 Pseudoleacina oligostropha (Reuss). – Pfeffer, p. 212. 869 Bulletin of Geosciences • Vol. 89, 4, 2014 non 1977 Pseudoleacina oligostropha (Reuss). – Moayedpour, p. 68, pl. 7, figs 2, 3. non 1999 Pseudoleacina (Paraglandina) oligostropha (Reuss). – Esu, p. 332. Material. – 2 specimens (NHMW 2013/0572/0041), 149 specimens (NHMW 1890/0013/0407, 1909/0001/0009). Dimensions. – Diameter: 4.5 mm, height: 13.6 mm (Fig. 9Q, R); diameter: 4.3 mm, height: 13.3 mm (Fig. 9S, T). Discussion. – Reuss in Reuss & Meyer (1849) considered the specimens from the Most Basin to be conspecific with Achatina sandbergeri Thomä, 1845 from the Chattian of Hochheim (Germany). This decision was also supported by the illustration of a Bohemian shell in which the growth lines and the spiral grooves are strongly overemphasised, suggesting an almost cancellate sculpture. Wenz (1917), however, pointed out that a separation of both species can be based on the nearly smooth shells of P. oligostropha. Boettger (1891) separated some rare narrower and more cylindrical shells as variation P. subcylindrata. Such morphologies are also found in our material but represent only slender morphotypes. Klika (1891) recognised the specimen described as Achatina oligostropha by Reuss (in Reuss & Meyer 1849) as a juvenile shell. Neglecting the rules of priority, he introduced Oleacina neglecta as new name for the adult shells of the very same species. Therefore, Oleacina neglecta Klika, 1891 is a subjective junior synonym of Achatina oligostropha Reuss. Esu (1999) assigned this species to the subgenus Paraglandina Pfeffer, 1930. The type species of this (sub)genus is the Late Oligocene Paraglandina confusa Pfeffer, 1930 by monotypy. It is characterised by its high spire, incised sutures and a blunt axial ribbing, bearing little in common with Pseudoleacina oligostropha. The specimens from the Early Miocene of Theobaldshof/Rhön (Germany) mentioned by Moayedpour (1977) differ in their stouter shape, the lower last whorl, the thin and narrow columellar lip and the deeper concavity of the columella. Occurrence. – Only known from Korozluky, Tuchořice, Lipno and Pyšná. Pseudoleacina producta (Reuss in Reuss & Meyer, 1849) Figure 9U–X 1849a A.[chatina] producta Rss.; Reuss in Reuss & Meyer, p. 11 (nomen nudum). *1849b A.[chatina] producta m.; Reuss in Reuss & Meyer, p. 32, pl. 3, fig. 12. 1861 Gl.[andina] producta Reuss. – Reuss, p. 70. 870 1875 Oleacina producta Reuss. – Sandberger, p. 444, pl. 24, figs 29, 29a. 1891 Oleacina producta Reuss. – Klika, p. 23, text-fig. 14. 1892 Oleacina producta Reuss. – Klika, p. 22, text-fig. 14. 1897 Oleacina producta Reuss var. emphysematica n. – Babor, p. 2. 1897 Oleacina subsulcosa (Thomae). – Babor, p. 17 (non Achatina subsulcosa Thomä, 1845). 1911 [Oleacina] producta Rss. – Kafka, p. 67. 1917 Poiretia (Pseudoleacina) producta (Reuss). – Wenz, p. 53. 1923 Poiretia (Pseudoleacina) producta producta (Reuss). – Wenz, p. 861 (cum syn.). 1923 Poiretia (Pseudoleacina) producta emphysematica (Babor). – Wenz, p. 862 (cum syn.). 1930 Pseudoleacina producta (Reuss). – Pfeffer, p. 213. non 2006 Pseudoleacina (Pseudoleacina) producta (Reuss), 1849 var. – Kókay, p. 85, pl. 32, figs 6, 7. Material. – 2 specimens (NHMW 2013/0572/0042), 160 specimens (NHMW 1890/0013/0407, 1909/0001/0010, 1909/0001/0098). Dimensions. – Diameter: 4.9 mm, height: 15.8 mm (Fig. 9U, V); diameter: 4.0 mm, height: 12.5 mm (Fig. 9W, X). Discussion. – A very characteristic and common species, with very high spire, strongly oblique sutures and strong axial ribs closely below the sutures. These ribs fade out quickly and grade into faint growth lines in the lower two thirds of the whorls. These features and the slender-bullet shaped outline allow a clear separation from Pseudoleacina oligostropha even for juvenile shells. Similarly, the record of Oleacina subsulcosa (Thomä, 1845) from Pyšná in a list of Babor (1897) might be based on strongly sculptured species of Pseudoleacina producta or P. oligostropha. The type specimen of Oleacina producta var. emphysematica Babor, 1897 does not justify any separation from Pseudoleacina producta. The Early Miocene Hungarian specimens, identified by Kókay (2006) as variety of Pseudoleacina producta, are clearly not conspecific with the Bohemian species. They differ in their much broader shells and the completely different type of axial sculpture. A further occurrence in the late Early Miocene of Vermes near Delsberg (= Delémont, Switzerland), described by Maillard (1891), is most probably not conspecific based on its stepped spire and higher aperture. Poiretia (Pseudoleacina) producta (Rss.) var. subcylindrica Wenz in Fischer & Wenz, 1915, from the Late Oligocene of Hochheim, was later transferred to P. (P.) sandbergeri by Wenz (1923). This species has a striking counterpart in the Recent Sardopoiretia emanueli Bodon, Nardi, Braccia & Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake Cianfanelli, 2010 from Sardinia, which differs in its more prominent and continuous axial sculpture and the lower protoconch and more convex first teleoconch whorl. Occurrence. – Only known from Tuchořice, Lipno and Pyšná. Superfamily Punctoidea Morse, 1864 Family Punctidae Morse, 1864 Genus Punctum Morse, 1864 Type species. – Helix minutissima Lea, 1841, by monotypy. Recent, North America. Punctum propygmaeum Andreae, 1904 Figure 12A–E *1904 Punctum propygmaeum sp. nov.; Andreae, p. 6, fig. 4. 1923 Punctum propygmaeum propygmaeum Andreae. – Wenz, p. 349. Finger (1998) differs clearly in its much coarser growth lines and the broader and more prominent striae on the protoconch. In addition, specimens from the Sarmatian of Austria, identified as P. parvulum by Harzhauser et al. (2008), have a much more prominent spiral sculpture on the teleoconch. Punctum oligocaenicum was described by Zinndorf (1901) from the Oligocene of Offenbach in Germany. This species differs from the Bohemian and Middle Miocene Punctum species in its down-bent aperture; in fact this species might not even belong to Punctum. Judging from the description and illustration given by Andreae (1904), the Bohemian shells agree in size and outline with those from Opole and therefore we tentatively assign the specimens to P. propygmaeum. Occurrence. – P. propygmaeum was originally described from the late Middle Miocene (MN7/8) from Opole in Poland. This tiny species has not been mentioned from the Most Basin so far and is recorded there only from Tuchořice. Family Discidae Thiele, 1931 (1866) Material. – 102 specimens (NHMW 2013/0572/0043). Genus Discus Fitzinger, 1833 Dimensions. – Diameter: 1.2 mm, height: 0.72 mm (Fig. 12A); diameter: 0.8 mm, height: 1.3 mm. Description. – Tiny shell with low conical spire and regularly convex whorls. The moderately convex protoconch has a diameter of 400 μm and displays a faintly malleated initial part; the following whorl is smooth on its adapical half and bears more or less developed spiral striae on its lower half, being most prominent close to the lower suture. The teleoconch is entirely covered by faint spiral threads which are crossed by weak, slightly sigmoidal growth lines. The spiral sculpture is weakest close to the upper suture and in the umbilical region. The umbilicus is deep and open; peristome thin with slightly widened inner lip. Discussion. – The Miocene representatives are usually identified as Punctum propygmaeum Andreae, 1904 and P. parvulum Gottschick, 1920 (Gottschick 1920a). Both taxa are known mainly from late Middle Miocene (Serravallian; late Badenian and Sarmatian) sections of Poland, Germany and Austria. SEM studies of Punctum parvulum from Steinheim in Finger (1998) document that this species is different from Late Miocene shells of the Vienna Basin, treated as P. propygmaeum by Lueger (1981). As comparable studies on topotypic material from Opole in Poland are missing so far, the relation between all these taxa remains unclear. The Middle Miocene (Sarmatian) Punctum parvulum Gottschick, 1920 from Steinheim in Germany sensu Type species. – Helix ruderata Hartmann, 1821; subsequent designation by Gray (1847). Recent, Europe. Discus bohemicus (Wenz in Fischer & Wenz, 1914) Figure 12I–K 1868 Helix multicostata Thom. – Reuss, p. 81, pl. 1, fig. 2 (non Helix multicostata Thomä, 1845). 1891 Patula multicostata Thomae. – Klika, p. 39, text-figs 31a–d (non Helix multicostata Thomä, 1845). 1892 Patula multicostata Thomae. – Klika, p. 38, text-figs 31a–d (non Helix multicostata Thomä, 1845). *1914 Pyramidula (Goniodiscus) bohemica sp. nov.; Wenz in Fischer & Wenz, p. 57. 1917 Pyramidula (Goniodiscus) bohemica Wenz. – Wenz, p. 55. 1923 Gonyodiscus (Gonyodiscus) bohemicus Wenz. – Wenz, p. 325 (cum syn.). 1964 Pyramidula (Gonyodiscus) euglypha (Reuss, 1849). – Čtyroký et al., p. 137, fig. 2 (non Helix euglypha Reuss in Reuss & Meyer, 1849). Material. – 2 specimens (NHMW 2013/0572/0044), 25 specimens (NHMW 1909/0001/0026). Dimensions. – Diameter: 4.2 mm, height: 2.1 mm (Fig. 12J); diameter: 4.1 mm, height: 1.58 mm (Fig. 12I, K). 871 Bulletin of Geosciences • Vol. 89, 4, 2014 Discussion. – This species was originally intermingled with the Oligocene Discus multicostatus (Thomä, 1845). As pointed out by Wenz in Fischer & Wenz (1914), the Bohemian species differs from the older one in its narrower coiling, deeper sutures and regularly convex whorls, lacking a keel or angulation. The text in Reuss (1868) is misguiding as he mentioned shells of up to 11 mm diameter and 5 mm height. Discus bohemicus, however, is always much smaller ranging around 4–4.5 mm in diameter. Maybe Reuss (1868) had a small specimen of the then undescribed Pleurodiscus falciferus (Boettger, 1870) at hand. The shell described as Pyramidula (Gonyodiscus) euglypha by Čtyroký et al. (1964) from the Kralupy drilling in the Most Basin is rather Discus bohemicus based on its large bulbous protoconch, the dense axial ribbing and the deeply incised sutures. Occurrence. – Only known from Tuchořice and the drilling at Kralupy. Discus euglyphus (Reuss in Reuss & Meyer, 1849) Figure 12L–N 1849a H.[elix] euglypha Rss.; Reuss in Reuss & Meyer, p. 11 (nomen nudum). *1849b H.[elix] euglypha m.; Reuss in Reuss & Meyer, p. 22, pl. 1, fig. 12. 1861 H.[elix] euglypha Reuss. – Reuss, p. 63. 1875 Patula euglypha Reuss. – Sandberger, p. 427 (pars). 1891 Patula euglypha Reuss. – Klika, p. 37, text-figs 29a–c. 1892 Patula euglypha Reuss. – Klika, p. 36, text-figs 29a–c. 1894 Patula Diezi sp. nov.; Clessin, p. 30. 1911 [Patula] euglypha Rss. – Kafka, p. 67. 1916 [Patula] euglypha Reuss. – Thuma, p. 83. 1917 Pyramidula (Gonyodiscus) euglypha (Reuss). – Wenz, p. 56. 1918 Pyramidula (Gonyodiscus) euglypha (Reuss). – Wenz, p. 8. 1923 Gonyodiscus (Gonyodiscus) diezi (Clessin). – Wenz, p. 327. 1923 Gonyodiscus (Gonyodiscus) euglyphus Reuss. – Wenz, p. 328. 1930 Goniodiscus euglyphus Reuss. – Pfeffer, p. 42. 1999 Discus (Discus) euglyphus (Reuss). – Esu, p. 332. non 1863 Helix (Patula) euglypha Reuss. – Sandberger, p. 389, pl. 35, figs 18, 18a (= Patula sandbergeri Clessin, 1894). non 1863 Patula (Charopa) euglypha Reuss. – Sandberger, p. 373, pl. 24, figs 3, 3a (= Patula sandbergeri Clessin, 1894). non 1864 Helix euglypha Reuss. – Deshayes, p. 820, pl. 9, figs 33–36 (= Patula sandbergeri Clessin, 1894). non 1964 Pyramidula (Gonyodiscus) euglypha (Reuss, 1849). – Čtyroký et al., p. 137, fig. 2 [= Discus bohemicus (Wenz in Fischer & Wenz, 1914)] non 1977 Discus (Discus) euglyphus (Reuss). – Moayedpour, p. 65, pl. 5, figs 5–9 (is probably D. rasseri sp. nov.). Material. – 2 specimens (NHMW 2013/0572/0045), 430 specimens (NHMW 1909/0001/0024). Dimensions. – Diameter: 4.55 mm, height: 1.9 mm (Fig. 12M); diameter: 4.3 mm, height: 1.8 mm (Fig. 12L, N). Discussion. – Differs from Discus bohemicus clearly in its lower shell, the less numerous but stronger axial ribs and the more or less developed angulation along the periphery. In addition, its protoconch is less convex and bears several somewhat irregular spiral grooves, which are deepest close to the lower suture. The strong and sigmoidal ribs on the base are a characteristic feature. When describing the Early/Middle Miocene MN5 faunas from Undorf, Clessin (1894) discussed also several Oligocene and Early Miocene taxa, including the Discidae from Tuchořice. He concluded that the Diez-collection contains a then undescribed species, which he named Patula Diezi. His description was mainly focussed on differences from Discus euglyphus and lacks an illustration and even data on the size of this new species. Clessin (1894) emphasised the low spire, wide umbilicus, a delicate and dense axial ribbing and the absence of an angulation. The rich material of the NHM collection documents that the angulation and spire height of D. euglyphus is quite variable and morphotypes as described by Clessin (1894) range well within D. euglyphus. Occurrence. – Known from Korozluky, Tuchořice and Lipno. Further occurrences from the Early Miocene of Donaurieden and Theobaldshof/Rhön in Germany need Figure 12. A–E – Punctum propygmaeum Andreae, 1904, NHMW 2013/0572/0043. • F–H – Manganellia alata (Klika, 1891), NHMW 2013/0572/0051. • I, J – Discus bohemicus (Wenz in Fischer & Wenz, 1914), NHMW 2013/0572/0044. • K – protoconch of Discus bohemicus, same specimen as I. • L, M – Discus euglyphus (Reuss in Reuss & Meyer, 1849), NHMW 2013/0572/0045. • N – protoconch of Discus euglyphus, same specimen as L. • O – Discus rasseri sp. nov., NHMW 2013/0572/0047 (paratype 1). • P – Discus rasseri sp. nov., NHMW 2013/0572/0046 (holotype). • Q – Discus rasseri sp. nov., NHMW 2013/0572/0048 (paratype 2). • R, S – Discus stenospira (Reuss in Reuss & Meyer, 1849), NHMW 2013/0572/0050. • T – protoconch of Discus stenospira, same specimen as R. 872 Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake D E F C A B G H I I L K P M J N R O Q S T 873 Bulletin of Geosciences • Vol. 89, 4, 2014 confirmation and are partly misidentifications (e.g. Moayedpour 1977). Discus rasseri Harzhauser, Neubauer & Georgopoulou sp. nov. Figure 12O–Q ? 1977 Discus (Discus) euglyphus (Reuss). – Moayedpour, p. 65, pl. 5, figs 5–9 (non Helix euglypha Reuss in Reuss & Meyer, 1849). Types. – Holotype (NHMW 2013/0572/0046): diameter: 2.95 mm, height: 1.55 mm (Fig. 12P); paratype 1 (NHMW 2013/0572/0047): diameter: 3.05 mm, height: 1.55 mm (Fig. 12O); paratype 2 (NHMW 2013/0572/0048): diameter: 2.95 mm, height: 1.55 mm (Fig. 12Q). Material. – 280 specimens (NHMW 1909/0001/0027). Etymology. – In honour of Michael W. Rasser, palaeontologist at the Stuttgart State Museum of Natural History. Diagnosis. – Small shell with striate protoconch, low conical spire, a distinct angulation in the upper third of the last whorl and a weakly convex lower part, causing a bowl-like outline. Prominent axial ribbing, which fades out quickly along the angulation and re-appears as narrow rim close to the umbilicus. Description. – Raised protoconch consisting of 1.5 convex whorls (850 μm diameter) with densely-spaced but very faint spiral threads, which are best developed in the lower half of the whorl. Teleoconch comprising 3.5 whorls; the spire displays some variability in height, ranging from low conical to nearly flat; sutures deeply incised. The first two teleoconch whorls bear regularly spaced, very prominent, prosocline-prosocyrt axial ribs, slightly narrower than the smooth interspaces. On the last whorl the interspaces become gradually wider and growth lines become stronger. A distinct angulation along the periphery separates the upper third of the last whorl from the weakly convex, rapidly contracting lower part. The axial ribs persist down to the angulation and are strongly prosocline; below the periphery they fade out quickly, passing into faint, slightly opisthocyrt growth lines and re-appear as distinct fold-like ribs again very close to the deep, sub-cylindrical umbilicus. Aperture semi-circular, moderately narrow with smooth peristome. Discussion. – This species was erroneously identified in the NHM collection as Janulus densestriatus (Klika, 1891) and most probably it was also intermingled with this species in the literature. Despite some similarities in sculpture 874 and shell outline, a placement within Janulus Lowe, 1852 is excluded based on the absence of any palatal teeth. Partly, this species was probably also mistaken for Discus euglyphus (e.g. specimens from the Early Miocene of Theobaldshof/Rhön in Moayedpour 1977). The Bohemian Discus rasseri species differs from D. euglyphus clearly in the strength and the higher position of the angulation, the narrower umbilicus and the weak axial ribs on the base. In addition, the semi-circular, strongly oblique aperture differs considerably from the wide and circular aperture of D. euglyphus. These features allow also a clear separation of the specimens of Moayedpour (1977) from D. euglyphus. The poor preservation, however, does not allow a decision whether these shells are conspecific with the Bohemian ones. Morphologically, the Bohemian species is very close to Patula gyrorbis Klein sensu Sandberger, 1875 (p. 454, pl. 21, figs 7–7b). A separation of D. rasseri from this Oligocene and Early Miocene German species is based on the much smaller size (about half of the diameter), the less numerous and wider spaced ribs on the upper shell surface and on the lower number of teleoconch whorls (3.5 versus 5.5). As pointed out by Manganelli et al. (2011), Helix gyrorbis Klein, 1846 (p. 72, pl. 1, figs 14a–c) is obviously not conspecific with Patula gyrorbis Klein sensu Sandberger, 1875. Moser et al. (2009) consider the latter species to represent the genus Janulus, but no information on internal denticles is available so far. Therefore, we tentatively treat it as a member of the Discidae. The Middle Miocene Discus costatus Gottschick sensu Finger, 1998 is also very similar to the Bohemian species. It differs in its smaller protoconch diameter (700 μm versus 800 μm), the higher spire, the less abrupt termination of the axial ribs, and the deeper position of the angulation. The original illustration of Discus costatus in Gottschick (1911, p. 501, pl. 7, figs 15a–c) shows a high-spired shell with prominent ribs on the base and a wide umbilicus, differing strongly from Discus rasseri. Occurrence. – Known from Tuchořice and probably also from the Early Miocene of Theobaldshof/Rhön (Germany). Discus stenospira (Reuss in Reuss & Meyer, 1849) Figure 12R–T 1849a Helix stenospira Rss.; Reuss in Reuss & Meyer, p. 12 (nomen nudum). *1849b H.[elix] stenospira m.; Reuss in Reuss & Meyer, p. 22, pl. 1, fig. 11. 1861 H.[elix] stenospira Reuss. – Reuss, p. 63. 1861 H.[elix] lunula Thom. – Reuss, p. 63 (non Helix lunula Thomä, 1845). Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake 1870a Helix (Patula) stenospira Rss. – Boettger, p. 287, pl. 13, figs 2a–c. 1891 Patula stenospira Reuss. – Klika, p. 38, text-fig. 30. 1892 Patula stenospira Reuss. – Klika, p. 37, text-fig. 30. 1911 [Patula] stenospira Rss. – Kafka, p. 67. 1914 Pyramidula (Goniodiscus) stenospira (Reuss). – Wenz in Fischer & Wenz, p. 58. 1915 Pyramidula (Goniodiscus) stenospira Reuss. – Fischer & Wenz, p. 45. 1916 [Patula] stenospira Reuss. – Thuma, p. 83. 1916 Pyramidula (Gonyodiscus) stenospira (Reuss). – Wenz, p. 167. 1917 Pyramidula (Gonyodiscus) stenospira (Reuss). – Wenz, p. 56. 1923 Gonyodiscus (Gonyodiscus) lunula stenospira (Reuss). – Wenz, p. 334 (cum syn.). 1977 Discus (D.) lunula stenospira (Reuss). – Moayedpour, p. 54. non 1891 Patula (Discus) stenospira Rss. sp. – Penecke, p. 359 [= Discus pleuradrus (Bourguignat, 1881)]. Material. – 2 specimens (NHMW 2013/0572/0050), 4 specimens (NHMW 1909/0001/0025). Dimensions. – Diameter: 3.9 mm, height: 2.2 mm (Fig. 12S); diameter: 3.9 mm (Fig. 12R, T). Description. – A very characteristic shell with strongly reduced sculpture consisting only of widely spaced, low and fold-like axial ribs, which are most prominent close to the upper suture. The microsculpture consists of dense and very delicate spiral furrows covering the entire teleoconch. These spiral furrows are even weaker on the otherwise smooth protoconch. Umbilicus deep and sub-cylindrical. Discussion. – The Bohemian species was often intermingled with the Early Miocene Discus lunula (Thomä, 1845) from the Mainz Basin. Later, Wenz (1923) and Moayedpour (1977) treated it as subspecies of D. lunula. Based on the description and illustration of this species by Sandberger (1858), both species can be distinguished by the very low and wide last whorl and the regular growth lines of D. lunula. Discus stenospira is superficially reminiscent of the Middle Miocene Lucilla subteres (Clessin, 1877). It differs from this and other Lucilla species in its much larger size, the less convex adapical part of the last whorl above the periphery and the less incised sutures. In addition, its last whorl is distinctly increasing in width but only weakly expanding in Lucilla. Occurrence. – Known from Korozluky, Tuchořice and Pyšná. Further occurrences are mentioned from the Early Miocene of Germany (Öpfingen, Hochheim, Ehingen, Theobaldshof/Rhön) by Wenz (1923) and Moayedpour B A C D Figure 13. Discus zagorseki sp. nov. • A – NHMW 2013/0572/0075 (paratype 1). • B –NHMW 2013/0572/0076 (paratype 2). • C – NHMW 2013/0572/0077 (holotype). • D – protoconch of A. (1977). Shells from the early Middle Miocene of the Rein Basin in Austria, identified as Discus stenospira by Penecke (1891), might rather represent Discus pleuradrus (Bourguignat, 1881; see Harzhauser et al. 2014 for discussion and further references). Discus zagorseki Harzhauser, Neubauer & Georgopoulou sp. nov. Figure 13A–D Types. – Holotype (NHMW 2013/0572/0077): diameter: 2.7 mm, height: 1.25 mm (Fig. 13C); paratype 1 (NHMW 2013/0572/0075): diameter: 2.75 mm, height: 1.25 mm (Fig. 13A, D); paratype 2 (NHMW 2013/0572/0076): diameter: 2.85 mm, height: 1.3 mm (Fig. 13B). Material. – 3 specimens. Etymology. – In honour of Kamil Zágoršek, palaeontologist at the National Museum in Prague. Diagnosis. – Small shell with striate protoconch, very low spire, prominent angulation below the spire and rapidly contracting, weakly convex base. Strong axial ribs on spire passing into weak ribs and growth lines along the periphery and base. Delicate spiral threads on base and prominent axial ribs in the circum-umbilical area. Wide and deep umbilicus; aperture narrow and oblique. Description. – A low discoidal shell consisting of 1.7 protoconch whorls and about 3 teleoconch whorls. Its moderately convex protoconch has a diameter of 700 μm and bears numerous spiral threads. The spire is very low conical with 875 Bulletin of Geosciences • Vol. 89, 4, 2014 moderately convex whorls bearing prominent axial ribs separated by slightly wider interspaces (55 ribs appear on the last whorl). The last whorl is characterised by a prominent angulation directly below the spire. The whorl is only weakly convex below the angulation and contracts rapidly towards the umbilicus. The ribs are prominent and strongly prosocline along the angulation and become much weaker and weakly opisthocyrt below. On the base, the weak ribs are sigmoidal and become prominent again in the circum-umbilical area. A delicate microsculpture of broad spiral threads, which are somewhat staggered along the growth lines, covers the base. Umbilicus wide and deep. Aperture narrow, oblique with subparallel margins and thin peristome. Discussion. – This species is superficially similar with Discus rasseri sp. nov. due to its near identical axial sculpture on the spire. Differences are the slightly smaller diameter, the lower spire, the more prominent circum-umbilical axial sculpture, and the pronounced angulation of Discus zagorseki. The transition from angulation into base is less convex and the angulation is in a more adapical position. Moreover, the protoconch of D. zagorseki is less convex but its spiral sculpture is better developed. The characteristic angulation of the periphery allows a clear separation from all other Miocene Discus species of the Most Basin. The early Miocene Discus neumaieri Schlickum, 1964 differs in its continuous axial ribs on the base (Binder 2004) and the convex periphery. The late Early to Middle Miocene Discus pleuradrus (Bourguignat, 1881), as illustrated by Böttcher et al. (2009), has a higher spire and its axial ribs continue along the flanks. shell and strongly prosocline along the keel/bulge. They become weak below the bulge and very prominent again in the umbilical area. Umbilicus moderately wide and deep; apertural margin thin. Discussion. – We place this genus in the family Discidae Thiele, 1931. Some species of Discus are slightly reminiscent of the new genus but all Miocene representatives of Discus of which SEM-data are available develop a completely different protoconch sculpture, being smooth or faintly striate (e.g. Harzhauser & Binder 2004, Böttcher et al. 2009, and species herein). The extant Discus perspectivus (Megerle von Mühlfeld, 1816) is superficially similar concerning the discoid and keeled shape but is distantly larger, develops a higher number of teleoconch whorls and has a very flat but still conical spire. The shells have a superficial similarity with Xerocrassa Monterosato, 1892. Although the type species of Xerocrassa (Helix seetzeni Pfeiffer, 1847) has little in common with the discoid fossil shells, several extant Xerocrassa species, such as Xerocrassa barceloi (Hidalgo, 1878), X. betulonensis (Bofill, 1897), and X. prietoi (Hidalgo, 1878) develop somewhat similar shells with comparable sculpture. However, these species are always distinctly larger compared to the tiny Manganellia species and lack the characteristic protoconch sculpture. Included species. – Two species are assigned to Manganellia. The type species M. alata (Klika, 1891) is an Early Miocene representative known only from the Most Basin in Bohemia. The second species is M. schneideri (Harzhauser & Neubauer in Harzhauser et al., 2014) from the early Middle Miocene of the Austrian Rein Basin. Occurrence. – Only known from Tuchořice. Occurrence. – Early and Middle Miocene of Central Europe. Genus Manganellia Harzhauser, Neubauer & Georgopoulou gen. nov. Type species. – Patula alata Klika, 1891. Early Miocene, Czech Republic. Etymology. – In honour of Giuseppe Manganelli, specialist for Neogene and Holocene terrestrial molluscs at the University of Siena (Italy). Diagnosis. – Minute, saucer-shaped shells with very low spire from which the large protoconch protrudes; the ca 800 μm wide protoconch consists of about 1.5 convex whorls with conspicuous crazing effect sculpture. Periphery formed by a keel or bulge in the upper part of the last whorl, which contracts quickly into the base via a more or less pronounced concavity. Sculpture consisting of prominent axial ribs, which are prosocyrt on the upper side of the 876 Manganellia alata (Klika, 1891) Figure 12F–I 1869a Helix (Patula) disculus A. Braun. – Slavík, p. 245, pl. 4, figs 3, 4 (non Helix disculus Sandberger, 1858). 1869b Helix (Patula) disculus A. Braun. – Slavík, p. 261, pl. 4, figs 3, 4 (non Helix disculus Sandberger, 1858). 1870a Helix (Patula) disculus A. Br. – Boettger, p. 288 (non Helix disculus Sandberger, 1858). *1891 Patula alata n.; Klika, p. 40, text-figs 33a–c. 1892 Patula alata Klika; Klika, p. 39, text-figs 33a–c. 1911 [Patula] alata Kl. – Kafka, p. 67. 1917 Pyramidula (Gonyodiscus) alata (Klika). – Wenz, p. 56. Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake 1923 Gonyodiscus (Gonyodiscus) alatus (Klika). – Wenz, p. 324. Material. – 2 specimens (NHMW 2013/0572/0051), 15 specimens (NHMW 1909/0001/0028). Dimensions. – Diameter: 3.3 mm, height: 1.2 mm (Fig. 12F, G); diameter: 3.3 mm, height: 1.25 mm (Fig. 12H, I). Discussion. – This rare, tiny disc-shaped shell is characterised by strongly raised axial ribs and a prominent bulge at the periphery, adjoined by a weak abapical concavity; its protoconch has a conspicuous crazing effect sculpture. The closest relative was recently described from the early Middle Miocene of Styria in Austria as “Discus” schneideri Harzhauser & Neubauer in Harzhauser et al. (2014). This species has an identical protoconch but differs in its larger number of axial ribs and the less discoidal outline. Moreover, it develops a keel at the periphery. The Oligocene “Discus” disculus (Braun in Walchner, 1851) is slightly reminiscent of the Bohemian shell concerning its discoidal outline, and strong axial sculpture but is clearly distinguished by its much wider umbilicus (see Sandberger 1858, p. 16, pl. 2, figs 10–10b and Sandberger 1875, p. 373, pl. 22, figs 12a–c). pelian of Dvérce, but from the context it seems very likely that Babor (1897) referred to material from Tuchořice as his entire paper considered only specimens from that locality. An additional occurrence at Tuchořice was mentioned by Klika (1891), who emphasised that he had seen only a single specimen in a local private collection without having the opportunity to compare the specimen with the Oligocene type material. In our opinion, it is unlikely that the Burdigalian specimens are conspecific with the much older Rupelian ones. Unfortunately, the specimens of Babor (1897) could not be detected in the collection of the National Museum in Prague. The generic status of this Early Miocene Janulus is confirmed by the presence of radial rows of palatal teeth described by Babor (1897) – but it remains unclear if this is only true for the Tuchořice specimens or also for the Rupelian ones as already discussed by Manganelli et al. (2011). Occurrence. – Janulus densestriatus is known from the Rupelian of Dvérce (Wärzen) and Dětaň (Czech Republic) (Mikuláš et al. 2003). Janulus sp. in only documented from Tuchořice. Family Pristilomatidae Cockerell, 1891 Genus Vitrea Fitzinger, 1833 Occurrence. – Only known from Tuchořice. Type species. – Glischrus (Helix) diaphana Studer, 1820; by monotypy. Recent, Europe. Superfamily Gastrodontoidea Tryon, 1866 Family Gastrodontidae Tryon, 1866 Vitrea cf. procrystallina (Andreae, 1902) Figure 14S–U Genus Janulus Lowe, 1852 Type species. – Helix (Janulus) calathus Lowe, 1852; by monotypy. Recent. Madeira. Janulus sp. 1891 Patula densestriata n.; Klika, p. 40 (pars). 1892 Patula densestriata Klika. – Klika, p. 39 (pars). 1897 Patula densestriata Klika. – Babor, p. 16 (non Patula densestriata Klika, 1891). 1923 Janulus densestriatus (Klika). – Wenz, p. 301 (pars). Material. – No material was available. Dimensions. – Unknown, but probably comparable to J. densestriata: diameter: 5.7 mm, height: 1.8 mm. Discussion. – Babor (1897) discussed a Janulus species, which he identified as Patula densestriata Klika, 1891. Janulus densestriatus was originally described from the Ru- cf. 1902b Hyalina (Vitrea) procrystallina sp. nov.; Andreae, p. 10, fig. 4. cf. 1923 Vitrea procrystallina procrystallina (Andreae). – Wenz, p. 293. cf. 1976 Vitrea procrystallina (Andreae). – Schlickum, p. 12, pl. 2, fig. 38. Material. – 2 specimens (NHMW 2013/0572/0052). Dimensions. – Diameter: 2.4 mm (Fig. 14S); diameter: 2.3 mm (Fig. 14T). Description. – Flat protoconch consisting of ca 0.75 nearly flat whorls with faint spiral sculpture. About two thinshelled teleoconch whorls form a tiny shell with low spire and nearly smooth surface except for faint spiral threads (more prominent than on the protoconch) and very delicate, orthocyrt growth lines. Smooth, regularly convex base with narrow umbilicus; even in SEM pictures the spiral sculpture is nearly invisible on the base. 877 Bulletin of Geosciences • Vol. 89, 4, 2014 Discussion. – The Bohemian species has a somewhat wider last whorl compared with the drawings in Andreae (1902b) and the internal cast in Schlickum (1976) but agrees well with the Late Miocene Vitrea p. steinheimensis Gottschick, 1920 sensu Lueger (1981, p. 42, pl. 5, fig. 7). The delicate spiral sculpture was also described by Lueger (1981). The status of the Middle Miocene V. p. procrystallina and V. p. steinheimensis and of the Late Miocene species described by Lueger (1981) is unclear as no adequate SEM illustrations are available so far. Therefore, we treat the Bohemian shells as “cf.” Occurrence. – This species was not recorded from the Most Basin so far. Vitrea p. procrystallina and V. p. steinheimensis are described from various Middle Miocene localities in Poland, Hungary and Germany (Wenz 1923, Kókay 2006) and from the Late Miocene of the Vienna Basin (Lueger 1981). (Fig. 14P); largest specimen: diameter: 3.8 mm, height: 1.7 mm. Discussion. – The spiral sculpture on the protoconch and the early teleoconch is only visible in SEM-images and consists of faint spiral grooves with interspaces of 7–10 μm width. The spiral furrows become most prominent on the last whorl. A delicate spiral thread coincides with the upper suture, forming a faint collar. These features, the low spire and the regularly convex base allow a clear separation from all other Oxychilus and Aegopinella species of the Most Basin. Schlickum & Strauch (1975) studied this species in their revision of Perpolita Baker, 1928 and documented that the Early Miocene shells from Germany, listed by Wenz (1923) as Zonitoides mendicus, represent a different species and that the Bohemian species belongs to Oxychilus. Genus Oxychilus Fitzinger, 1833 Occurrence. – This species is known only from the Burdigalian of Tuchořice; the poorly preserved, juvenile shells from the much younger Late Badenian and Sarmatian of Hungary described by Kókay (2006) are not conspecific with O. mendicus; they are higher and the sutures are distinctly more incised. Type species. – Helix cellaria Müller, 1774; original designation. Recent, Europe. Subfamily Godwiniinae Cooke, 1921 Family Oxychilidae Hesse in Geyer, 1927 (1879) Subfamily Oxychilinae Hesse in Geyer, 1927 (1879) Oxychilus mendicus (Slavík, 1869) Figure 14O–R *1869a Helix (Hyalina) mendica sp. nov.; Slavík, p. 247, pl. 4, figs 7, 8. 1869b Helix (Hyalina) mendica sp. nov.; Slavík, p. 262, pl. 4, figs 7, 8. 1891 Hyalinia mendica Slavík. – Klika, p. 31, text-fig. 23. 1892 Hyalinia mendica Slavík. – Klika, p. 31, text-fig. 23. 1911 [Hyalinia] mendica Sl. – Kafka, p. 67. 1917 Hyalinia (Hyalinia) mendica Slavik. – Wenz, p. 54. 1923 Zonitoides (Zonitoides) mendicus (Slavik). – Wenz, p. 296 (pars; Bohemian occurrences only). 1975 Helix (Hyalinia) mendica Slavik. – Schlickum & Strauch, p. 41. non 1902b Hyalina (Polita) mendica Slavic [sic]. – Andreae, p. 8 (= Perpolita wenzi Schlickum & Strauch, 1975). non 1918 Hyalina (Polita) mendica Slavik. – Wenz, p. 7 (= Perpolita wenzi Schlickum & Strauch, 1975). non 2006 Nesovitrea (Perpolita) mendica Slavik, 1869. – Kókay, p. 76, pl. 29, figs 1–5. Material. – 2 specimens (NHMW 2013/0572/0053), 20 specimens (NHMW 1909/0001/0018). Dimensions. – Diameter: 3.6 mm, height: 1.9 mm 878 Genus Aegopinella Lindholm, 1927 Type species. – Helix pura Alder, 1830; original designation. Recent, Great Britain. Aegopinella denudata (Reuss in Reuss & Meyer, 1849) Figure 14D–G 1849a H. [elix] denudata Rss.; Reuss in Reuss & Meyer, p. 11 (nomen nudum). *1849b H. [elix] denudata m.; Reuss in Reuss & Meyer, p. 21, pl. 1, fig. 9. 1861 H.[elix] denudata Reuss. – Reuss, p. 64. 1875 Hyalinia denudata Reuss. – Sandberger, p. 441, pl. 24, figs 23–23b. 1891 Hyalinia denudata Reuss. – Klika, p. 28, text-fig. 19. 1892 Hyalinia denudata Reuss. – Klika, p. 28, text-fig. 19. 1897 Hyalina denudata Reuss var. sculpta n. – Babor, p. 2. 1911 Hyalinia denudata Rss. – Kafka, p. 67. 1915 Hyalina (Hyalina) denudata Reuss. – Fischer & Wenz, p. 45, pl. 2, fig. 3. 1916 Hyalinia denudata Reuss. – Thuma, p. 83. 1917 Hyalinia (Hyalinia) denudata (Reuss). – Wenz, p. 54. 1923 Oxychilus (?Oxychilus) denudatum denudatum (Reuss). – Wenz, p. 274 (cum syn.). Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake 1923 Oxychilus (?Oxychilus) denudatum sculptum (Babor). – Wenz, p. 274. 1930 Hyalinia (Polita) denudata Reuss. – Pfeffer, p. 36. 1977 Retinella (Retinella) denudata (Reuss). – Moayedpour, p. 66. 2004 Aegopinella denudata (Reuss, 1849). – Binder, p. 197, pl. 3, fig. 4. ? 2006 Aegopinella denudata (Reuss), 1849 var. – Kókay, p. 77, pl. 30, figs 1, 2. Material. – 2 specimens (NHMW 2013/0572/0055), 246 specimens (NHMW 1847/0032/0092, 1890/0013/0395, 1909/0001/0015, 1909/0001/0099). Dimensions. – Diameter: 10 mm, height: 4.2 mm (Fig. 14D–F); largest specimen: diameter: 14 mm, height: 4.5 mm. Discussion. – The generic status of this species changed considerably in the literature. Wenz (1923) listed it as Oxychilus and Moayedpour (1977) considered it to be a Retinella [the identifications were done by Schlickum as mentioned by Moayedpour in the introduction], whilst Binder (2004) described it as Aegopinella. This view is followed herein as the depressed shell outline, the delicate microscopical spiral striation, the wide and elongate-ovoid aperture and the rapidly expanding last whorl correspond much better to Aegopinella. Babor (1897) separated a shell with prominent axial ribs and spiral threads on the base as “var. sculpta”. The very fragmentary specimen is still present in the Prague collection (NM-PM-P 458) and does not support a taxonomic separation. Occurrence. – Known from Korozluky, Tuchořice and Lipno; additional occurrences are mentioned from the Early Miocene of Theobaldshof/Rhön (Germany; middlelate Burdigalian) and Oberdorf (Austria; mid-Burdigalian). It is also reported from the Hungarian Early Miocene (Kókay 2006), although the larger size and the additional teleoconch whorl of the specimens from the Somlóvásárhely drilling differ from typical A. denudata (Kókay 2006). Aegopinella vetusta (Klika, 1891) Figure 15A–C *1891 1892 1911 1923 Hyalinia vetusta n.; Klika, p. 31, text-figs 22a–c. Hyalinia vetusta Klika. – Klika, p. 30, text-figs 22a–c. [Hyalinia] vetusta Kl. – Kafka, p. 67. Oxychilus (? Oxychilus) vetustum (Klika). – Wenz, p. 287 (cum syn.). ? 1977 Retinella (Retinella) cf. vetusta (Klika). – Moayedpour, p. 66, pl. 6, figs 4–6. Material. – One specimen in the Prague collection (NM-PM-P 741). Dimensions. – Diameter: 10 mm, height: 4.2 mm. Description. – Depressed solid shell with nearly completely flat spire and moderately incised suture and wide last whorl with regularly convex periphery. Prominent growth lines cover the spire and the periphery, where they are distinctly prosocline. Base smooth and glossy; umbilicus wide and open. Discussion. – According to Hartmut Nordsieck (written comm. 2014), this species does not represent the genus Retinella Fischer in Shuttleworth, 1877, as suggested by Moayedpour (1977), but might rather represent an Aegopinella. A specimen from the Burdigalian of Theobaldshof/Rhön described by Moayedpour (1977) is very similar to the species described by Klika (1891) concerning shape, umbilical features and incised sutures. The poor preservation of the shell surface, however, does not allow a clear identification. Moreover, the less convex upper part of the periphery of the shell from Germany differs from the regularly convex periphery of A. vetusta. Occurrence. – Only known from Pyšná. Superfamily Zonitoidea Mörch, 1864 Family Zonitidae Mörch, 1864 Discussion. – Several rather poorly defined genera have been described for Oligocene and Miocene Zonitidae by Pfeffer (1930) and Lueger (1981). Consequently, species are variously treated as Archaeozonites, Miozonites and Pontaegopis (see Binder 2004, Moser et al. 2009, Harzhauser et al. 2014). Herein, we provisionally accept Miozonites Pfeffer, 1930 mainly because the Most Basin species Helix algiroides is its type species. Genus Miozonites Pfeffer, 1930 Type species. – Helix algiroides Reuss in Reuss & Meyer, 1849; subsequent designation by Wenz & Zilch (1959). Early Miocene, Czech Republic. Miozonites algiroides (Reuss in Reuss & Meyer, 1849) Figure 14H–M 1849a Helix algiroides Rss.; Reuss in Reuss & Meyer, p. 11 (nomen nudum). 1849a H.[elix] Haidingeri Rss.; Reuss in Reuss & Meyer, p. 11 (nomen nudum). 879 Bulletin of Geosciences • Vol. 89, 4, 2014 1849a H. [elix] semiplana Rss.; Reuss in Reuss & Meyer, p. 11 (nomen nudum). *1849b H.[elix] algiroides m.; Reuss in Reuss & Meyer, p. 19, pl. 1, fig. 5. 1849b H.[elix] Haidingeri m.; Reuss in Reuss & Meyer, p. 19, pl. 1, fig. 6. 1849b H. [elix] semiplana m.; Reuss in Reuss & Meyer, p. 20, pl. 1, figs 7, 8. 1861 H.[elix] algiroides Reuss. – Reuss, p. 63. 1861 H.[elix] Haidingeri Reuss. – Reuss, p. 63. 1861 H.[elix] semiplana Reuss. – Reuss, p. 63. 1869a Zonites algiroides Reuss. – Slavík, p. 238. 1869a Zonites semiplanus Reuss. – Slavík, p. 239. 1870a Helix (Campylaea Beck) semiplana Rss. – Boettger, p. 290. 1875 Archaeozonites semiplanus Reuss. – Sandberger, p. 442, pl. 24, figs 25–25b. 1875 Archaeozonites Haidingeri Reuss. – Sandberger, p. 404, 443, pl. 24, figs 26–26b. 1890 Archaeozonites Haidingeri Reuss. – Klika, p. 43, fig. 14 1a–c. 1890 Archaeozonites Haidingeri var. Reussi. – Klika, p. 43, fig. 14 2a, b. 1891 Archaeozonites Haidingeri Reuss. – Klika, p. 25, text-figs 17a–c. 1891 [Archaeozonites Haidingeri] var. Reussi Klika. – Klika, p. 27, text-figs 18a, b. 1892 Archaeozonites Haidingeri Reuss. – Klika, p. 25, text-figs 17a–c. 1892 [Archaeozonites Haidingeri] var. Reussi Klika. – Klika, p. 27, text-figs 18a, b. 1911 [Archaeozonites] Haidingeri Rss. – Kafka, p. 67. 1911 [Archaeozonites Haidingeri] var. Reussi Kl. – Kafka, p. 67. 1914 Zonites (Aegopis) algiroides (Reuss). – Wenz in Fischer & Wenz, p. 48. 1916 Archaeozonites Haidingeri Reuss. – Thuma, p. 83. 1916 [Archaeozonites] Haidingeri var. Reussi Klika. – Thuma, p. 83. 1917 Zonites (Aegopis) algiroides (Reuss). – Wenz, p. 54. 1923 Zonites (Aegopis) algiroides (Reuss). – Wenz, p. 248 (cum syn.). 1930 Miozonites algiroides (Reuss) var. Reussi (Klika). – Pfeffer, p. 177. 1932 Zonites (Aegopis) algiroides (Reuss). – Wenz, p. 19. 1933 Archaeozonites (Miozonites) algiroides (Reuss). – Wenz, p. 8. 2006 Aegopis algiroides (Reuss), 1849. – Kókay, p. 76, pl. 29, fig. 10. 2010 Helix. – Dvořák et al., p. 58, unnumbered plate, fig. 4. Material. – 2 specimens (NHMW 2013/0572/0056), 58 specimens (NHMW 1855/0034/0032, 1890/0008/0394, 1909/0001/0013, 1909/0001/0014, 1909/0001/0017, 1909/0001/0022). Dimensions. – Diameter: 26 mm, height: 17.5 mm (Fig. 14H–J); diameter: 28 mm, height: 16.2 mm (Fig. 14K–M). Discussion. – This species is one of the largest and most characteristic gastropods in the Tuchořice collections, attaining a diameter of more than 40 mm. It is thin shelled and displays considerable variability concerning spire height and the more or less distinct angulation of the last whorl. Characteristic features are the deep, perspectivic, moderately wide umbilicus, the thin peristome and the sculpture consisting of densely spaced, thin but prominent prosocline to prosocyrt ribs on the upper part of the whorls. These ribs are of varying strength and may rarely be bifurcate and become very weak or fade out on the base. Few subsutural spiral grooves cross the axial ribs in most specimens. The allometric growth with a rapidly expanding last whorl allows a clear distinction between juvenile and adult shells. All these factors resulted in the introduction of different species and varietal names for the same species (e.g. Helix semiplana, Helix haidingeri, Archaeozonites haidingeri var. reussi; M. semiplanus is a juvenile stage of M. algiroides; see also Fischer & Wenz 1914). Occurrence. – Known from Korozluky, Tuchořice and Lipno; a tiny fragment from the Lower Miocene of the Somlóvásárhely drilling in Hungary described by Kókay (2006) might represent a “subplanus”-morph, but the preservation does not allow a clear identification; the irregular axial ribs, however, are fully comparable with M. algiroides and differ from the late Early/Middle Miocene M. costatus (Sandberger, 1875). Additional occurrences are listed by Wenz (1923, 1932, 1933) from the Late Oligocene to Early Miocene of Germany (Hochheim-Flörsheim, Eggingen, Hauchenberg/Missen) and France (Son-Saucats). Figure 14. A–C – Lyrodiscus ihli (Klika, 1891), NHMW 2013/0572/0054. • D–F – Aegopinella denudata (Reuss in Reuss & Meyer, 1849), NHMW 2013/0572/0055. • G – Aegopinella denudata (Reuss in Reuss & Meyer, 1849), NHMW 2013/0572/0055. • H–M – Miozonites algiroides (Reuss in Reuss & Meyer, 1849), NHMW 2013/0572/0056. • O, P – Oxychilus mendicus (Slavík, 1869), NHMW 2013/0572/0053. • Q, R – microsculpture of Oxychilus mendicus (Slavík, 1869), NHMW 2013/0572/0053. • S–U – Vitrea cf. procrystallina (Andreae, 1902), NHMW 2013/0572/0052. • V, W – Phenacolimax intermedius (Reuss in Reuss & Meyer, 1849), NHMW 2013/0572/0058. • X, Y – Phenacolimax intermedius (Reuss in Reuss & Meyer, 1849), NHMW 2013/0572/0058. 880 Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake A D H K B E I L F C J M G O Q V T S P U R W X Y 881 Bulletin of Geosciences • Vol. 89, 4, 2014 Genus Lyrodiscus Pilsbry, 1893 Type species. – Helix circumsessa Shuttleworth, 1852; original designation. Recent. Canary Islands. Lyrodiscus ihli (Klika, 1891) comb. nov. Figure 14A–C *1891 1892 1911 1917 1923 Hyalinia ihli n.; Klika, p. 29, text-fig. 20. Hyalinia ihli Klika. – Klika, p. 29, text-fig. 20. [Hyalinia] Ihli Kl. – Kafka, p. 67. Hyalinia (Hyalinia) ihli Klika. – Wenz, p. 54. Oxychilus ihli (Klika). – Wenz, p. 277 (cum syn.). Material. – 2 specimens (NHMW 2013/0572/0054), 15 specimens (NHMW 1901/0001/0016). Dimensions. – Diameter: 8 mm, height: 4.3 mm (Fig. 14A–C); largest specimen: diameter: 9.2 mm, height: 4.7 mm. Discussion. – Hartmut Nordsieck (written comm. 2014), suggested a placement in Lyrodiscus (Riedeliella) Schlickum & Strauch, 1975, based on the sculpture of the base of this species. Own investigations supported this view. The base of well preserved specimens bears very delicate spiral threads and few widely spaced spirals consisting of short discontinuous and somewhat irregularly staggered parts. This pattern is characteristic for several species of the genus Lyrodiscus as illustrated by Rousseau & Puisséger (1990). As shown by Alonso et al. (2013), Riedeliella is a junior synonym of Lyrodiscus. Only three specimens of this species were known to Klika (1891) when describing this shell, which was probably frequently intermingled with Aegopinella denudata. The richer material of the NHMW collection supports the description of Klika (1891) and suggests only minor variability in shape. Thus, the conical spire, the coarse growth lines (which are most prominent at the upper suture), the smaller size, the narrower last whorl and the less incised sutures of L. ihli allow a clear separation from A. denudata. The placement in the family Zonitidae follows Alonso et al. (2013). [= Limacus flavus (Linnaeus, 1758)]; by monotypy. Recent, Europe (anthropogenic in Australia). Limacus crassitesta (Reuss, 1868) Figure 15D *1868 1869a 1891 1892 ? 1902b ? 1904 ? 1915 1911 1917 1923 1964 2009 non 1923 Limax crassitesta Rss.; Reuss, p. 81, pl. 1, fig. 1. Limax crassitesta Reuss. – Slavík, p. 239. Sansania crassitesta Reuss. – Klika, p. 20 (pars). Sansania crassitesta Reuss. – Klika, p. 20 (pars). Sansania crassitesta (Reuss). – Andreae, p. 7. Sansania crassitesta (Rss). – Andreae, p. 3. Sansania crassitesta (Reuss). – Fischer & Wenz, p. 44. Saussania [sic] crassitesta Rss. – Kafka, p. 67. Sansania crassitesta (Reuss). – Wenz, p. 53. Limax crassitesta (Reuss). – Wenz, p. 309 (pars). Milax crassitesta (Reuss, 1868). – Čtyroký et al., p. 137, fig. 1. “Limax” crassitesta Reuss 1868. – Moser et al., p. 49. Limax cf. crassitesta Reuss. – Wenz, p. 310 (cum syn.). Material. – One specimen (NHMW 2013/0572/0057), 4 specimens in the Prague collection (NM-PM-P 496). Dimensions. – Diameter: 2.3 mm, length: 3.3 mm (NM-PM-P 496). Discussion. – The generic placement of this species in Limacus follows Moser et al. (2009). The broad and robust specimens in the NHMW and Prague collections agree well with the type illustrated by Reuss (1868). Only the specimen illustrated by Klika (1891) displays a more slender outline and might represent a second species. It is doubtful that the many records from the Late Oligocene and Early Miocene of Germany and the Middle Miocene of Poland listed by Wenz (1923) all represent the same species. Late Miocene specimens, listed by Wenz (1923) as “cf. crassitesta”, are most probably not conspecific with the Early Miocene species. Occurrence. – Known from Tuchořice and the Kralupy drillings. Occurrence. – Only known from Tuchořice. Family Vitrinidae Fitzinger, 1833 Superfamily Limacoidea Lamarck, 1801 Family Limacidae Lamarck, 1801 Genus Limacus Lehmann, 1864 Type species. – Limacus breckworthianus Lehmann, 1864 882 Genus Phenacolimax Stabile, 1859 Type species. – Helicolimax major Férussac, 1807; subsequent designation by Fischer in Paulucci (1878). Recent, Europe. Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake Phenacolimax intermedius (Reuss in Reuss & Meyer, 1849) Figure 14V–Y 1849a Vitrina intermedia Rss.; Reuss in Reuss & Meyer, p. 11 (nomen nudum). *1849b V.[itrina] intermedia m.; Reuss in Reuss & Meyer, p. 18, pl. 1, fig. 4. ? 1858 Vitrina intermedia Reuss. – Sandberger, p. 12, pl. 5, figs 19–19c. 1875 Vitrina intermedia Reuss. – Sandberger, p. 426, pl. 24, figs 27–27c. 1861 V.[itrina] intermedia m. – Reuss, p. 62. 1891 Vitrina intermedia Reuss. – Klika, p. 23, text-figs 15/1. 1892 Vitrina intermedia Reuss. – Klika, p. 23, text-figs 15/1. 1911 Vitrina intermedia Rss. – Kafka, p. 67. 1916 Vitrina intermedia Reuss. – Thuma, p. 83. 1917 Vitrina (Phenacolimax) intermedia Reuss. – Wenz, p. 53. 1923 Vitrina intermedia intermedia Reuss. – Wenz, p. 216. ? 1977 ?Vitrina cf. intermedia Reuss. – Moayedpour, p. 65, pl. 5, figs 10, 11. non 1981 Semilimax intermedius (Reuss). – Lueger, p. 41, pl. 5, figs 1–3. non 2006 Semilimax intermedia (Reuss), 1849. – Kókay, p. 75, pl. 28, fig. 5. non 2004 Semilimax cf. intermedia (Reuss, 1852). – Harzhauser & Binder, p. 25, pl. 9, fig. 7. Kókay (2006) differ in their higher and (partly) much thicker shells, the Late Miocene Pannonian shells as described by Lueger (1981) and Harzhauser & Binder (2004) lack the microsculpture and their last whorls are expanding less quickly. Similarly, occurrences from Opole (Poland) mentioned by Andreae (1902b) need verification. These records should probably better be compared with Phenacolimax suevicus (Sandberger) sensu Schütt (1967). Occurrence. – Known from Korozluky, Tuchořice and Pyšná; a further occurrence from the Burdigalian of Theobaldshof/Rhön was mentioned by Moayedpour (1977). Phenacolimax crassitesta (Andreae, 1902) 1891 Vitrina intermedia Reuss. – Klika, p. 23, text-fig. 15/2 (non Vitrina intermedia Reuss in Reuss & Meyer, 1849). 1892 Vitrina intermedia Reuss. – Klika, p. 23, text-fig. 15/2 (non Vitrina intermedia Reuss in Reuss & Meyer, 1849). *1902b Vitrina (Semilimax) intermedia var. crassitesta n. nom.; Andreae, p. 7. 1923 Vitrina intermedia crassitesta Andreae. – Wenz, p. 217 (cum syn.). non 2006 Semilimax intermedia crassitesta (Andreae), 1902. – Kókay, p. 75, pl. 28, figs 6, 7. Material. – No material was available. Material. – 2 specimens (NHMW 2013/0572/0058), 7 specimens (NHMW 1909/0001/0011). Dimensions. – Diameter: 7.0 mm, height: 3.9 mm (Fig. 14X, Y); diameter: 4.8 mm, height: 3.2 mm (Fig. 14V, W). Discussion. – The generic status of most fossil vitrinids is doubtful (Nordsieck 2013b). Moser et al. (2009) placed the closely related Vitrina suevica Sandberger, 1872 in Phenacolimax Stabile, 1859, which is followed herein. Fragments of this fragile species are not rare in the studied collections. Only more or less complete and adult specimens reveal the characteristic ear-shaped outline due to the rapidly widening last whorl. The protoconch bears coarse and irregularly arranged pits, which become spirally arranged on its lower side (if viewed from the umbilical side) and continue as weaker, spirally arranged pits on the entire base. The apical part of the glossy shell is smooth aside from faint growth lines. Weak and discontinuous grooves appear on the base close to and parallel to the columellar lip. This seemingly long-lived species was also mentioned from Middle and Late Miocene strata. Whilst the Middle Miocene Sarmatian specimens from Hungary described by Dimensions (after Klika 1891). – Diameter: 6.6 mm, height: 5.2 mm. Discussion. – Andreae (1902b) refers to the thick shelled specimens from Tuchořice described by Klika (1891) as Vitrina intermedia. Whether the Middle Miocene specimens from Opole (Poland) represent the same species, however, remains dubious. Similarly, the identity of the shells from the Sarmatian of Hungary described by Kókay (2006) is dubious. The neritid-like shape and the rather straight columella suggest that these shells are not conspecific with P. crassitesta. Occurrence. – Only known from Tuchořice. Superfamily Helicoidea Rafinesque, 1815 Family Hygromiidae Tryon, 1866 Genus Leucochroopsis O. Boettger, 1908 Type species. – Leucochroa (Leucochroopsis) emmerichi O. Boettger, 1908; by monotypy. Miocene, Germany. 883 Bulletin of Geosciences • Vol. 89, 4, 2014 Leucochroopsis apicalis (Reuss, 1861) Figure 15E–K *1861 H.[elix] apicalis Reus [sic]; Reuss, p. 64, pl. 1, fig. 1. 1875 Helix (Fruticicola) leptoloma var. apicalis Reuss. – Sandberger, p. 380, pl. 24, figs 7–7c. 1870a Helix (Fruticicola) leptoloma A. Br. – Boettger, p. 289 (non Helix leptoloma Sandberger, 1858). 1891 Helix (Trichia) apicalis Reuss. – Klika, p. 55, text-figs 51a–c. 1892 Helix (Trichia) apicalis Reuss. – Klika, p. 35, text-figs 51a–c. 1897 Helix (Carthusiana Kobelt) oxyspira sp. nov.; Babor, p. 4, fig. 1. 1911 [Helix] (Trichia) apicalis Rss. – Kafka, p. 68. 1915 Trichopsis apicalis Reuss. – Fischer & Wenz, p. 47. 1916 [Helix] (Trichia) apicalis Reuss. – Thuma, p. 83. 1917 Hygromia (Trichopsis) apicalis (Reuss). – Wenz, p. 56. 1917 Hygromia (Monacha) oxyspira (Baber) [sic]. – Wenz, p. 57. 1923 Trichia (Leucochroopsis) apicalis apicalis (Reuss). – Wenz, p. 422 (cum syn.). 1923 Hygromia (Monacha) oxyspira (Babor). – Wenz, p. 415. 1930 Leucochroopsis apicalis apicalis (Reuss). – Pfeffer, p. 116. 1930 Zenobiella (Monachoides) [oxyspira (Babor)]. – Wenz, p. 3028. 1933 Trichia (Leucochroopsis) apicalis (Reuss). – Wenz, p. 8. ? 1977 Leucochroopsis apicalis (Reuss). – Moayedpour, p. 69, pl. 7, figs 6–8. 1980 [Monacha] oxyspira (Babor, 1897). – Richardson, p. 215. 1980 [Leucochroopsis] apicalis (Reuss, 1860). – Richardson, p. 455. usually develops a very weak angulation in its upper third. The peristome is thin, the outer lip is only slightly expanded and the columellar lip is gently reflected, covering parts of the narrow and deep umbilicus. A single specimen from Tuchořice was described by Babor (1897) as Helix oxyspira. It shows a bulbous last whorl and rather high conical spire; the spire appears small compared to the large last whorl due to the rapidly increasing width of the whorls. The raised protoconch consists of 1.3 strongly convex whorls. Shell smooth except for irregular, blunt growth lines, which are most prominent close to the incised sutures. Deep and open umbilicus with smooth circum-umbilical area bearing a weak ridge, which increases in strengths towards the aperture; thin peristome; the semi-circular aperture is expanding along its umbilical part, forming a slight concavity coinciding with the termination of the umbilical ridge and covering the umbilicus partly. Discussion. – Helix oxyspira Babor, 1897 seems to be a pathologic specimen of Leucochroopsis apicalis (Reuss, 1861). It has an identical protoconch and the characteristic ratio between pointed spire and wide last whorl. The strange umbilical ridge represents the only difference. As this feature is also reflected in the indention of the peristome, we consider this a healed injury. Therefore, we consider this species as subjective junior synonym of Leucochroopsis apicalis. Occurrence. – Known from Korozluky, Tuchořice, Lipno and Pyšná; the specimen from the Early Miocene of Theobaldshof/Rhön illustrated by Moayedpour (1977) is smaller and the last whorl is only weakly expanding but might represent a subadult shell of L. apicalis. An additional Early Miocene occurrence is mentioned by Wenz (1933) from Hauchenberg at Missen in Germany. Genus Pseudomonacha Pfeffer, 1930 Material. – 2 specimens (NHMW 2013/0572/0059), 138 specimens (NHMW 1890/0013/0398, 1909/0001/0043), 1 specimen (NM-PM-P 454, = Helix oxyspira Babor, 1897). Dimensions. – Diameter: 11 mm, height: 7 mm (Fig. 15E–H); diameter: 13 mm, height: 9.1 mm (Fig. 15I–K = holotype of Helix oxyspira Babor, 1897). Description. – A characteristic species with pointed conical spire and somewhat allometric, broad last whorl, which Type species. – Helix punctigera Thomä, 1845; subsequent designation by Nordsieck (1986). Early Miocene, Germany. Pseudomonacha zippei (Reuss in Reuss & Meyer, 1849) Figure 15L–S 1849a H. [elix] Zippei Rss.; Reuss in Reuss & Meyer, p. 11 (nomen nudum). Figure 15. A–C – Aegopinella vetusta (Klika, 1891), NM-PM-P 741. • D – Limacus crassitesta (Reuss, 1868), NM-PM-P 496. • E–H – Leucochroopsis apicalis (Reuss, 1861), NHMW 2013/0572/0059. • I–K – Leucochroopsis apicalis (Reuss, 1861), NM-PM-P 454 (= holotype of Helix oxyspira Babor, 1897). • L–S – Pseudomonacha zippei (Reuss in Reuss & Meyer, 1849), NHMW 2013/0572/0060. • T–Za – Pseudomonacha homalospira (Reuss, 1861), NHMW 2013/0572/0061. 884 Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake C A B E F G L M N O I H J P T X U Y Q R S D V W K Z Za 885 Bulletin of Geosciences • Vol. 89, 4, 2014 *1849b H.[elix] Zippei m.; Reuss in Reuss & Meyer, p. 24, pl. 2, figs 5, 6. 1861 H.[elix] Zippei Reuss. – Reuss, p. 64. 1875 Helix (Fruticicola) Zippei Reuss. – Sandberger, p. 428, pl. 24, figs 4–4c. 1891 Helix zippei Reuss. – Klika, p. 51, text-figs 46a, b. 1892 Helix zippei Reuss. – Klika, p. 49, text-figs 46a, b. 1909 [Monacha] zippei Reuss. – Boettger, p. 15. 1911 [Helix (Fruticicola)] Zippei Rss. – Kafka, p. 68. 1916 [Helix (Trigonostoma)] Zippei Reuss. – Thuma, p. 83. 1917 Hygromia (Monacha) zippei (Reuss). – Wenz, p. 57. 1923 Monacha (Monacha) zippei (Reuss). – Wenz, p. 418 (cum syn.). 1930 Pseudomonacha Zippei (Reuss). – Pfeffer, p. 112. 1930 Zenobiella (Monachoides) [zippei (Reuss)]. – Wenz, p. 3028. 1980 [Monachoides] zippei (Reuss, 1849). – Richardson, p. 464. 1999 Pseudomonacha zippei (Reuss). – Esu, p. 332. Material. – 2 specimens (NHMW 2013/0572/0060), 148 specimens (NHMW 1864/0012/725, 1890/0013/0398, 1909/0001/0040, 1909/0001/0041, 1909/0001/0042). Dimensions. – Diameter: 17 mm, height: 13 mm (Fig. 15L–O); diameter: 14.5 mm, height: 8.6 mm (Fig. 15P–S). Discussion. – This medium-sized species is characterised by its conical spire, the slightly angulated last whorl and the deeply incised furrow behind the flaring peristome. There is complete confusion between this species and the morphologically similar Pseudomonacha homalospira (Reuss, 1861) in many collections. Usually, the conical spire of P. zippei differs considerably from the much lower spire of P. homalospira. Nevertheless, there is some overlap between low-spired P. zippei and high-spired P. homalospira. Both species can be distinguished immediately based on the microsculpture: P. zippei develops a very delicate and densely spaced microsculpture of tiny papillae. This sculpture is not always fully preserved but usually visible on the last whorl; in addition, many specimens bear weak spiral grooves on the base. Pseudomonacha homalospira lacks any spiral grooves and bears coarser knob-like papillae accompanied by axially elongated pits. These structures override a faint velvet-like microsculpture formed by tiny papillae comparable to those of P. zippei. Moreover, some shells of P. zippei bear a single, narrow spiral band coinciding with the angulation of the last whorl; no comparable band is seen in P. homalospira. Both species, however, develop a dense pattern of papillae on their protoconchs. 886 Occurrence. – Only known from Korozluky, Tuchořice, Lipno and Pyšná. Pseudomonacha homalospira (Reuss, 1861) Figure 15T–Za *1861 H.[elix] homalospira Reuss; Reuss, p. 65, pl. 1, fig. 3. 1875 Helix homalospira Reuss. – Sandberger, p. 429, pl. 24, figs 6–6c. 1891 Helix homalospira Reuss. – Klika, p. 53, text-figs 48a–c. 1892 Helix homalospira Reuss. – Klika, p. 51, text-figs 48a–c. 1915 Fruticicola homalospira Reuss. – Fischer & Wenz, p. 47. 1909 [Monacha] homalospira Reuss. – Boettger, p. 15. 1911 [Helix] homalospira Rss. – Kafka, p. 68. 1916 [Helix (Trigonostoma)] homalospira Reuss. – Thuma, p. 83. 1917 Hygromia (Monacha) homalospira (Reuss). – Wenz, p. 57. 1923 Monacha (Monacha) homalospira (Reuss). – Wenz, p. 413 (cum syn.). 1930 Pseudomonacha homalospira (Reuss). – Pfeffer, p. 113. 1930 Zenobiella (Monachoides) [homalospira (Reuss)]. – Wenz, p. 3028. 1977 Monacha homalospira (Reuss). – Moayedpour, p. 54. 1980 [Monachoides] homalospira (Reuss, 1860). – Richardson, p. 461. non? 2006 Perforatella (Monachoides) homalospira (Reuss), 1860. – Kókay, p. 86, pl. 33, fig. 1. Material. – 2 specimens (NHMW 2013/0572/0061), 74 specimens (NHMW 1909/0001/0045, 1909/0001/0101), 7 specimens from Lipno (NHMW 1864/0012/0731). Dimensions. – Diameter: 18 mm, height: 8.5 mm (Fig. 15T–W); diameter: 15.5 mm, height: 8.2 mm (Fig. 15X–Za). Discussion. – See above (Pseudomonacha zippei). Occurrence. – Known from Korozluky, Tuchořice and Lipno; a further occurrence is mentioned without description by Moayedpour (1977) from the Early Miocene of Theobaldshof/Rhön (Germany). The Early Miocene specimen from the Somlóvásárhely drilling in Hungary, illustrated by Kókay (2006) seems to represent a different species based on the much coarser papillae and the prominent growth lines. Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake Family Helicodontidae Kobelt, 1904 Genus Praeoestophorella Pfeffer, 1930 Type species. – Helix phacodes Thomä, 1845, designation herein. Late Oligocene, Germany. Praeoestophorella petersi (Reuss in Reuss & Meyer, 1849) comb. nov. Figure 16A–F 1849a H.[elix] Petersi Rss.; Reuss in Reuss & Meyer, p. 11 (nomen nudum). *1849b H.[elix] Petersi m.; Reuss in Reuss & Meyer, p. 23, pl. 2, fig. 3. 1861 H.[elix] phacodes Thom. – Reuss, p. 68 (non Helix phacodes Thomä, 1845). 1891 Helix (Gonostoma) phacodes Thomae. – Klika, p. 45, text-figs 38a–c (non Helix phacodes Thomä, 1845). 1892 Helix (Gonostoma) phacodes Thomae. – Klika, p. 44, text-figs 38a–c (non Helix phacodes Thomä, 1845). 1897 Helix (Caracollina Ehrenberg) phacodes Thomae var. grossa n. – Babor, p. 3. 1911 Helix (Gonostoma) phacodes Th. – Kafka, p. 67 (non Helix phacodes Thomä, 1845). 1917 Helicodonta (Caracollina) phacodes (Thomae). – Wenz, p. 60 (non Helix phacodes Thomä, 1845). 1930 Praeoestophorella tuchoricensis nov.; Pfeffer, p. 127. 1980 [Caracollina] tuchoricensis Pfeffer, 1929. – Richardson, p. 426. Material. – 2 specimens (NHMW 2013/0572/0062), 29 specimens (NHMW 1890/0013/0397, 1909/0001/0100, 1909/0001/00033). Dimensions. – Diameter: 9.8 mm, height: 3.9 mm (Fig. 16A–C); diameter: 9.9 mm, height: 4.5 (Fig. 16D–F). Discussion. – This species was originally described as Helix Petersi Reuss in Reuss & Meyer, 1849. Later, Reuss (1861) doubted the validity of his species and suggested that is a junior synonym of Helix phacodes Thomä, 1845. This view was accepted by following authors up to Wenz (1923), until Pfeffer (1930) recognised the differences between both taxa and described the species from Tuchořice as Praeoestophorella tuchoricensis, erroneously overlooking the older available name petersi [and also the available name grossa of Babor (1897)]. Wenz (1917, 1923) treated this species as Caracollina and Richardson (1980) considered Praeoestophorella as synonym of Caracollina. Aside from a superficial similar- ity concerning outline and sculpture, the completely covered umbilicus allows a clear separation from Caracollina. The distinguishing features from the Oligocene P. phacodes, such as the much more pronounced keel in P. petersi and its lower discoidal shape and differences in the ribbing, were discussed in detail by Pfeffer (1930) and confirmed by us by comparison with material from Hochheim. Pfeffer (1930, p. 278) introduced Praeoestophorella as new genus without designating a type species. He listed Praeoestophorella phacodes (Thomä) and P. tuchoricensis Pfeffer as sole species of this genus. In the same monograph, Pfeffer (1930, p. 276) erected Pachycarocollina as new genus and listed only Pachycarocollina lapicidella (Thomä) as species within this genus, which thus is automatically the type species by monotypy (ICZN 1999, Article 68.3). Unfortunately, Nordsieck (1986) overlooked this decision and defined Helix phacodes Thomä as type species of Pachycarocollina. Herein, we designate Helix phacodes Thomä, 1845 as type species of Praeoestophorella Pfeffer, 1930, which corresponds to the original intention of Pfeffer (1930). The specimen described as Helix (Caracollina) phacodes var. grossa by Babor (1897) is a high spired morph with rather convex spire. After examination of the type in the Prague collection, we consider it an aberrant or unusual specimen of Praeoestophorella petersi. Occurrence. – Only known from Tuchořice and Pyšná; it might represent the direct descendant of the Oligocene to Aquitanian P. phacodes. Genus Protodrepanostoma Germain, 1929 Type species. – Helix (Polygyra) plioauriculata Sacco, 1889; original designation. Pliocene, Italy. Protodrepanostoma involuta nordsiecki Falkner, 1986 Figure 16 O–P 1849a H.[elix] involuta Thom. – Reuss in Reuss & Meyer, p. 11 (non Helix involuta Thomä, 1845). 1849b H.[elix] involuta Thom. – Reuss in Reuss & Meyer, p. 28, pl. 3, fig. 3 (non Helix involuta Thomä, 1845). 1861 H.[elix] involuta Thom. – Reuss, p. 68 (non Helix involuta Thomä, 1845). 1891 [Helix (Trigonostoma Fitz.) involuta Thomae] var. minor. – Klika, p. 47, text-figs 39a, b. 1892 [Helix (Trigonostoma Fitz.) involuta Thomae] var. minor. – Klika, p. 45, text-figs 39a, b. 1911 [Helix] (Trigonostoma) involuta Th. var. minor. – Kafka, p. 67. 1916 [Helix] (Trigonostoma) involuta Thomae var. minor Klika. – Thuma, p. 83. 887 Bulletin of Geosciences • Vol. 89, 4, 2014 1917 Helicodonta (Helicodonta) involuta (Thomae). – Wenz, p. 59 (non Helix involuta Thomä, 1845). 1930 Helicodonta involuta involuta (Thomae). – Pfeffer, p. 131 (pars) (non Helix involuta Thomä, 1845). *1986 Protodrepanostoma nordsiecki sp. nov.; Falkner, p. 121, pl. 17, figs 9, 10. 1999 Protodrepanostoma nordsiecki Falkner. – Esu, p. 332. 2000b “P.” involutum – nordsiecki. – Manganelli & Giusti, p. 352. Material. – 2 specimens (NHMW 2013/0572/0063), 85 specimens (NHMW 1890/0013/0403, 1909/0001/0034). Dimensions. – Diameter: 6.8 mm, height: 3.1 mm; diameter: 5.5 mm, height: 2.9 mm (Fig. 16P). Discussion. – Falkner (1986) placed this species in the genus Protodrepanostoma Germain, 1929 with the Pliocene type species Helix (Polygyra) plioauriculata Sacco, 1889. Manganelli & Giusti (2000b) suggested that this genus is a synonym of Drepanostoma Porro, 1836 and excluded P. nordsiecki from that genus as understood by them. Indeed, the wide aperture of the Miocene species differs significantly from D. helenae Manganelli & Giusti, 2000b and D. plioauriculatum Sacco, 1889, which develop a prominent parietal lamella. The presence of this lamella, however, is not a distinguishing feature and is missing in related species such as Helicodonta obvoluta sensu Ciangherotti et al. 2007 and may be absent or present within P. bernardii (Michaud, 1862) (H. Nordsieck, written comm. 2014). We cannot follow the statement of Manganelli & Giusti (2000b) that the Most Basin species is “practically indistinguishable” from the Oligocene and Early Miocene P. i. involuta (Thomä, 1845). In his precise description, Falkner (1986) listed a number of features, which allow a separation of both taxa at least on the subspecies level (e.g. the larger diameter of the papillae in P. i. nordsiecki). Occurrence. – Only known from Korozluky, Tuchořice and Lipno. Protodrepanostoma hecklei (Klika, 1891) Figure 16G–N *1891 [Helix (Trigonostoma Fitz.) involuta Thomae] var. hecklei; Klika, p. 47, text-figs 40a, b. 1892 [Helix (Trigonostoma Fitz.) involuta Thomae] var. hecklei. – Klika, p. 45, text-figs 40a, b. 1911 Helix (Trigonostoma) involuta Th. var. Hecklei; Kafka, p. 67. 1986 [Helicodonta] hecklei. – Falkner, p. 123, pl. 17, fig. 11. 1999 Helicodonta hecklei (Klika). – Esu, p. 332. Material. – 2 specimens (NHMW 2013/0572/0064), 4 specimens (NHMW 1909/0001/0102, 1909/0001/0035/). Dimensions. – Diameter: 8.8 mm, height: 3.4 mm (Fig. 16G–J); diameter: 7.8 mm, height: 3.4 mm (Fig. 16K–N). Discussion. – This species is readily distinguished from the somewhat similar Protodrepanostoma involuta nordsiecki by its larger size and the much weaker sculpture of early teleoconch whorls. According to Nordsieck (2013b, written comm. 2014), this group, including taxa such as P. bernardii (Michaud, 1862) and P. obvoluta sensu Ciangherotti et al. 2007, belongs to Protodrepanostoma and not to Helicodonta as proposed by Manganelli & Giusti (2000b) based on the apertural features (see Nordsieck 2013b, figs 6–9). Occurrence. – Only known from Tuchořice. Family Elonidae Gittenberger, 1979 Genus Metacampylaea Pilsbry, 1894 Type species. – Helix Rahtii Thomä, 1845; original designation. Oligocene, Germany. Metacampylaea papillifera (Klika, 1891) Figure 16Q–R *1891 Helix (Geotrochus?) papillifera n.; Klika, p. 64, text-figs 60a–c. 1892 Helix (Geotrochus?) papillifera Klika. – Klika, p. 62, text-figs 60a–c. 1911 Helix (Geotrochus) papillifera. – Kafka, p. 68. 1917 Metacampylaea papillifera (Klika). – Wenz, p. 57. 1921 [Metacampylaea (Metacampylaea)] papillifera (Klika). – Boettger & Wenz, p. 17. 1923 Metacampylaea (Metacampylaea) papillifera (Klika). – Wenz, p. 482 (cum syn.). Figure 16. A–F – Praeoestophorella petersi (Reuss in Reuss & Meyer, 1849), NHMW 2013/0572/0062. • G–J, K–N – Protodrepanostoma hecklei (Klika, 1891), NHMW 2013/0572/0064. • O, P – Protodrepanostoma involuta nordsiecki Falkner, 1986, NHMW 2013/0572/0063. • Q, R – Metacampylaea papillifera (Klika, 1891), NM-PM-P 432 (holotype); R – details of the microsculpture. • S–X – Pseudochloritis robusta (Reuss in Reuss & Meyer, 1849), NHMW 2013/0572/0065. • Y–Za – Apula prominens (Babor, 1897), NM-PM-P 707 (holotype). 888 Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake D A K G B E H C F L I M J N P O V S Q Y T W Z R U X Za 889 Bulletin of Geosciences • Vol. 89, 4, 2014 1933 Metacampylaea (Metacampylaea) papillifera (Klika). – Wenz, p. 8. 1980 [Metacampylaea (Galactochiloides)] papillifera (Klika, 1891). – Richardson, p. 58. Material. – One specimen in the Prague collection (NM-PM-P 432). Dimensions. – Diameter: 23.5 mm, height: 10.7 mm. Description. – Depressed, obtuse-carinate shell characterised by its sculpture consisting of large and wide-spaced papillae, arranged in strongly opisthocline rows. The entire shell is covered by a crazing effect microsculpture, consisting of densely spaced, slightly elongate to wrinkled papillae. All these elements are predominated by blunt, rather irregular, prosocline growth lines. A distinct angulation appears on the last whorl, which terminates in a strongly reflected peristome. Umbilicus covered. Unfortunately, the holotype is glued to a collection label and we did not dare to remove it. Therefore, we have to refer to Klika (1891) for accurate illustrations of the umbilical and apertural views. 1849a H.[elix] trichophora Rss.; Reuss in Reuss & Meyer and Meyer, p. 11 (nomen nudum). *1849b H.[elix] robusta m.; Reuss in Reuss & Meyer, p. 25, pl. 2, fig. 7. 1849b H.[elix] trichophora m.; Reuss in Reuss & Meyer, p. 26, pl. 2, fig. 8. 1861 H.[elix] robusta Reuss. – Reuss, p. 67. 1861 H.[elix] trichophora Reuss. – Reuss, p. 68. 1875 Helix (Hemicycla) robusta Reuss. – Sandberger, p. 431, pl. 24, figs 10–10b. 1891 Helix (Chlorites) robusta Reuss. – Klika, p. 60, text-figs 56a, b. 1892 Helix (Chlorites) robusta Reuss. – Klika, p. 58, text-figs 56a, b. 1897 Helix (Trachia Albers) Ihliana sp. nov.; Babor, p. 6, text-fig. 2. 1911 [Helix] (Chloritis) robusta Rss. – Kafka, p. 68. 1917 Tropidomphalus (Pseudochloritis) robustum (Reuss). – Wenz, p. 58. 1917 Tropidomphalus (Tropidomphalus) ihlianus (Babor). – Wenz, p. 58. 1923 Tropidomphalus (Pseudochloritis) ihlianus (Babor). – Wenz, p. 510 (cum syn.). 1923 Tropidomphalus (Pseudochloritis) robustus (Reuss). – Wenz, p. 517. 1930 Pseudochloritis robusta Reuss. – Pfeffer, p. 74. 1980 [Tropidomphalus (Pseudochloritis)] robustus (Reuss, 1849). – Richardson, p. 60. ? 2006 Tropidomphalus (Pseudochloritis) robustus (Reuss), 1852. – Kókay, p. 89, pl. 34, figs 5, 6, text-fig. 14. 2008 Pseudochloritis robusta (Reuss, 1849). – Binder, p. 171, pl. 1, figs 3a–c, 4a–c, 5a–c. Discussion. – An extremely rare species, which is known so far only from a single specimen described by Klika (1891; 20 additional specimens were mentioned by Klika from the private collection of O. Boettger). As discussed by Klika (1891) and Wenz (1917), this species seems to be closely related with the Oligocene Metacampylaea rahtii (Thomä, 1845), which is the type species of Metacampylaea Pilsbry, 1894. Therefore, the decision of Richardson (1980) to place this species in Galactochiloides cannot be followed, because this (sub)genus was introduced by Wenz (1919) for Metacampylaea-like species without keel. The weaker keel, lower spire and fully covered umbilicus allow a separation from M. rahtii. Material. – 2 specimens (NHMW 2013/0572/0065), 50 specimens (NHMW 2005z0059/0004, 2005z0059/0006, 2005z0059/0007, 2005z0059/0008), one specimen (NM-PM-P 453 = Helix ihliana Babor, 1897). Occurrence. – Known from Tuchořice; Wenz (1933) mentions the species also from the Early Miocene of Kustersberg (Germany) and Wirtatobel (Austria). Dimensions. – Diameter: 28 mm, height: 17.5 mm (Fig. 16S–U); diameter: 28 mm, height: 16 mm (Fig. 16V–X). Genus Pseudochloritis C. Boettger, 1909 Type species. – Helix incrassata Klein, 1853; original designation. Miocene, Germany. Pseudochloritis robusta (Reuss in Reuss & Meyer, 1849) Figure 16S–X 1849a H.[elix] robusta Rss.; Reuss in Reuss & Meyer, p. 11 (nomen nudum). 890 Discussion. – The species was discussed in detail by Binder (2008); the generally depressed spherical shell shows some variability concerning spire height and the covering of the deep umbilicus by the broad and reflected inner lip. Occurrence. – Common species in the Most Basin, known from Tuchořice and Lipno. Poorly preserved fragments described by Kókay (2006) from the Lower Miocene of the Somlóvásárhely drilling in Hungary might also represent this species. Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake Genus Klikia Pilsbry, 1895 Type species. – Helix osculum Thomä, 1845; original designation. Miocene, Germany. Klikia labiata (Klika, 1891) Figure 17G–L 1849a H.[elix] osculum Thom. – Reuss in Reuss & Meyer, p. 11 (non Helix osculum Thomä, 1845). 1849b H.[elix] osculum Thom. – Reuss in Reuss & Meyer, p. 27, pl. 3, fig. 2 (non Helix osculum Thomä, 1845). 1861 H.[elix] osculum Thom. – Reuss, p. 64 (non Helix osculum Thomä, 1845). 1891 Helix (Trigonostoma) osculum Thomae. – Klika, p. 47, text-figs 41a–c (non Helix osculum Thomä, 1845). *1891 [Helix (Trigonostoma) osculum Thomae] mut. labiata n.; Klika, p. 48, text-figs 42a–c. 1891 [Helix (Trigonostoma) osculum Thomae] mut. tenuis n.; Klika, p. 49, text-figs 43a, b. 1892 Helix (Trigonostoma) osculum Thomae. – Klika, p. 46, text-figs 41a–c (non Helix osculum Thomä, 1845). 1892 [Helix (Trigonostoma) osculum Thomae] mut. labiata Klika. – Klika, p. 47, text-figs 42a–c. 1892 [Helix (Trigonostoma) osculum Thomae] mut. tenuis Klika. – Klika, p. 47, text-figs 43a, b. 1911 [Helix] osculum mut. labiata. – Kafka, p. 67. 1911 Helix osculum mut. tenuis. – Kafka, p. 67. 1911 Klikia labiata Klika. – Wenz, p. 96, pl. 4, figs 31–35. 1916 [Helix (Trigonostoma)] osculum Thomae var. tenuis Klika. – Thuma, p. 83. 1916 Klikia (Klikia) osculum tenuis Klikia [sic]. – Wenz, p. 58. 1917 Klikia (Klikia) osculum (Thomae). – Wenz, p. 47 (non Helix osculum Thomä, 1845). 1917 Klikia (Klikia) labiata (Klika). – Wenz, p. 58. 1923 Klikia (Klikia) labiata (Klika). – Wenz, p. 545 (cum syn.). 1923 Klikia (Klikia) osculum tenuis (Klika). – Wenz, p. 552 (cum syn.). 1930 [Klikia] labiata. – Pfeffer, p. 86. 1980 [Klikia] labiata (Klika, 1891). – Richardson, p. 56. Material. – 4 specimens (NHMW 2013/0572/0066), 443 specimens (NHMW 1847/0032/0091, 1864/0012/0729, 1909/0001/0036, 1909/0001/00037, 1909/0001/00038, 1909/0001/00052, 1909/0001/0103). Dimensions. – Diameter: 9 mm, height: 6 mm (Fig. 17J–L); diameter: 9.5 mm, height: 6.5 mm (Fig. 17G–I). Discussion. – This very frequent species has a solid, depressed, globose shell with low spire, strongly convex last whorl and base, and a typical, strongly thickened, reflected, collar-like peristome with a glossy welldefined parietal callus. There is some minor variability in sphericity and especially in the thickness of the peristome. Such shells with reflected but rather thin outer lip have been treated as “var. tenuis” by Klika (1891) and Wenz (1923). In our opinion, they just represent late subadult stages. This species is most likely a direct descendant of the slightly older Oligocene and Aquitanian Klikia osculum (Thomä, 1845), which lacks the thickened peristome. Occurrence. – Only known from Korozluky, Tuchořice and Lipno. Genus Apula C. Boettger, 1909 Type species. – Helix devexa Reuss, 1861; original designation. Early Miocene, Czech Republic. Apula devexa (Reuss, 1861) Figure 17A–F *1861 1869a 1891 1891 1897 1875 1911 1911 1912 1916 1923 1933 1930 1960 1980 non 1915 non 2006 H.[elix] devexa Reuss; Reuss, p. 65, pl. 1, fig. 4. Helix (Fruticicola) devexa Reuss. – Slavík, p. 242. Helix devexa Reuss. – Klika, p. 50, text-figs 45a–c. Helix devexa Reuss. – Klika, p. 49, text-figs 45a–c. Helix devexa Reuss f. applanata n.; Babor, p. 3. Helix (Fruticicola) devexa Reuss. – Sandberger, p. 429, pl. 24, figs 5–5c. Helix (Fruticicola) devexa Rss. – Kafka, p. 68. Klikia devexa Rss. – Wenz, p. 94, pl. 4, figs 26–28. Klikia devexa Reuss. – Boettger, p. 129. Klikia (Apula) devexa (Reuss). – Wenz, p. 59. Klikia (Apula) devexa (Reuss). – Wenz, p. 537 (pars). Klikia (Apula) devexa (Reuss). – Wenz, p. 9. Klikia (Apula) devexa (Reuss). – Pfeffer, p. 87. Klikia (Apula) devexa (Reuss). – Zilch in Wenz & Zilch, p. 707, fig. 2459. [Klikia (Ampula [sic])] devexa (Reuss, 1860). – Richardson, p. 58. Klikia (Apula) devexa (Rss). – Fischer & Wenz, p. 49. Klikia (Apula) cf. devexa (Reuss) var., 1860. – Kókay, p. 92, pl. 35, figs 11–15. Material. – 2 specimens (NHMW 2013/0572/0067), 58 specimens (NHMW 1890/0013/0399, 1909/0001/0039), 3 specimens in the Prague collection (NHM-PM-P 879-881, = Helix devexa applanata Babor, 1897). 891 Bulletin of Geosciences • Vol. 89, 4, 2014 Dimensions. – Diameter: 10 mm, height: 5.2 mm (Fig. 17D–F); diameter: 11 mm, height: 6.3 mm (Fig. 17A–C). Family Helicidae Rafinesque, 1815 Discussion. – All available specimens are quite homogenous concerning size, shape and umbilicus. The umbilicus is nearly completely covered or reduced to a narrow puncture. The microsculpture consists of a dense pattern of delicate papillae. The specimens separated by Babor (1897) as “applanata” do not justify any taxonomic separation. Type species. – Helix obtusecarinata Sandberger, 1858; original designation. Early Miocene, Czech Republic. Occurrence. – Known from Tuchořice and Lipno. Wenz & Fischer (1915) mentioned this species also from the Early Miocene of Theobaldshof/Rhön (Germany). Based on the wider umbilicus of these specimens, Moayedpour (1977) doubted this identification. Similarly, the Early Miocene specimens from the Hungarian Somlóvásárhely drilling, tentatively assigned to Apula devexa by Kókay (2006), differ distinctly in their much wider umbilicus. An Early Miocene occurrence from Wirtatobel (Austria) is mentioned by Wenz (1933). Apula prominens (Babor, 1897) species inquirenda Figure 16Y–Za 1897 Helix devexa Reuss f. prominens n. – Babor, p. 4. Material. – One specimen in the Prague collection (NM-PM-P 707). Dimensions. – Diameter: 8.5 mm, height: 5.8 mm. Discussion. – A single specimen was described by Babor (1897) as variation of Apula devexa. Later, Wenz (1923) listed this taxon as synonym of Apula devexa (Reuss, 1861). At first sight, it differs clearly from that common species in its high spire, the convex whorls, the comparatively narrower last whorl (in apical view) and the wider umbilicus. In addition it has a large and bulbous protoconch consisting of ca 0.75 whorls. A closer look at the protoconch, however, documents a pathologic development with an injury early in ontogeny. Its microsculpture is identical with that of A. devexa and the peristome is also very similar. Therefore, the status of this species remains unclear unless further specimens are found. Occurrence. – Known from Tuchořice. Genus Creneatachea Zilch in Wenz & Zilch, 1960 Creneatachea obtusecarinata (Sandberger, 1858) Figure 17U–Zb 1849a Helix Rahtii Thom. – Reuss in Reuss & Meyer, p. 11 (non Helix Rahtii Thomä, 1845). 1849b H.[elix] Rahtii Thomae. – Reuss in Reuss & Meyer, p. 24, pl. 2, figs 1–2 (non Helix Rahtii Thomä, 1845). *1858 Helix obtusecarinata; Sandberger, p. 25. 1861 H.[elix] obtusecarinata Sandb. – Reuss, p. 66. 1875 Helix obtusecarinata Sandberger. – Sandberger, p. 430, pl. 24, figs 9–9c. 1870a Helix (Iberus Montf.) obtuse carinata [sic] Sandb. – Boettger, p. 291. 1891 Helix (Trichia) perfecta n.; Klika, p. 56, text-figs 52a, b. 1891 Helix (Geotrochus?) obtusecarinata Sandberger. – Klika, p. 62, text-figs 57a, b. 1891 [Helix (Geotrochus?) obtusecarinata Sandberger] forma minima n. – Klika, p. 63, text-fig. 58. 1891 [Helix (Geotrochus?) obtusecarinata Sandberger] var. obesula n. – Klika, p. 63, text-figs 59a–c. 1892 Helix (Geotrochus?) obtusecarinata Sandberger. – Klika, p. 60, text-figs 57a, b. 1892 [Helix (Geotrochus?) obtusecarinata Sandberger] forma minima Klika. – Klika, p. 61, text-fig. 58. 1892 [Helix (Geotrochus?) obtusecarinata Sandberger] var. obesula Klika. – Klika, p. 61, text-figs 59a–c. 1909 [Tachea] obtusecarinata Sdbg. – Boettger, p. 18. 1911 [Helix] (Trichia) perfecta Kl. – Kafka, p. 68. 1911 [Helix] (Geotrochus? Hemicyclus) obtusecarinata Sndb. – Kafka, p. 68. 1911 [Helix (Geotrochus? Hemicyclus) obtusecarinata Sndb.] var. minima Kl. – Kafka, p. 68. 1912 Helix (Tachea?) obtusecarinata Sdb. – Frankenberger, p. 73. 1916 [Helix] (Geotrochus?) obtusecarinata Sandberger. – Thuma, p. 83. 1917 Cepaea obtusecarinata (Sandberger). – Wenz, p. 61. 1917 Cepaea subsoluta (Sandberger). – Wenz, p. 60 [non Helix (Crena) subsoluta Sandberger, 1858]. 1918 Cepaea obtusecarinata (Sandberger). – Wenz, p. 17. Figure 17. A–F – Apula devexa (Reuss, 1861), NHMW 2013/0572/0067. • G–L – Klikia labiata (Klika, 1891), NHMW 2013/0572/0066. • M–P, S–T – Megalotachea macrocheila (Reuss in Reuss & Meyer, 1849), NHMW 2013/0572/0069. • Q, R – Megalotachea macrocheila (Reuss in Reuss & Meyer, 1849), MB.Ga.12215. • U–Zb – Creneatachea obtusecarinata (Sandberger, 1858), NHMW 2013/0572/0068. 892 Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake A B H E C M K F I U Q O S P T L Y V Z W Za R N J G D X Zb 893 Bulletin of Geosciences • Vol. 89, 4, 2014 1923 Cepaea obtusecarinata (Sandberger). – Wenz, p. 648 (cum syn.). 1933 Cepaea obtusecarinata (Sandberger). – Wenz, p. 9. 1930 Crena obtusecarinata Sandb. – Pfeffer, p. 189. 1960 Cepaea (Creneatachea) obtusecarinata (Sandberger). – Zilch in Wenz & Zilch, p. 717, text-fig. 2491. 1980 [Cepaea (Creneatachea)] obtusecarinata (Sandberger, 1858). – Richardson, p. 286. Material. – 2 specimens (NHMW 2013/0572/0068), 149 specimens (NHMW 1843/0040/0002, 1847/0032/0089, 1890/0013/0396, 1909/0001/0053, 1909/0001/0049). Dimensions. – Diameter: 16 mm, height: 9.9 mm (Fig. 17U–X); diameter: 21 mm, height: 12 mm (Fig. 17Y–Zb). Discussion. – A very conspicuous lenticular and nonumbilicate shell – usually with very prominent carina – which cannot be confused with any other species. The broad, raised and flattened basal lip is another characteristic feature of this species. Along with the typical morphotypes appear rare specimens with tiny umbilical chink (mainly subadult shells) and ones, which lack the carina resulting in a rather depressed spherical outline. Some variation can also be stated concerning the spire height and the strength of the growth lines. Colour patterns are rarely preserved and seem to consist only of two narrow brown bands on the base. Occurrence. – Known from Korozluky, Tuchořice and Lipno; an additional occurrence from the Lower Miocene of Ulm (Germany) and Wirtatobel (Austria) is mentioned by Wenz (1923, 1933). Genus Megalotachea Pfeffer, 1930 Type species. – Helix turonensis Deshayes, 1831; subsequent designation by Truc (1971). Early Miocene, France. Megalotachea macrocheila (Reuss in Reuss & Meyer, 1849) nov. comb. Figure 17M–T 1849a Helix oxystoma Thom. – Reuss in Reuss & Meyer, p. 11 (non Helix oxystoma Thomä, 1845). 1849a H.[elix] macrocheila Rss.; Reuss in Reuss & Meyer, p. 12 (nomen nudum). *1849b H.[elix] macrocheila m.; Reuss in Reuss & Meyer, p. 26, pl. 3, fig. 1. 1849b H.[elix] rostrata A. Braun. – Reuss in Reuss & Meyer, p. 27, pl. 2, fig. 9 (non “Helix rostrata Braun”). 1861 H.[elix] deflexa A. Braun. – Reuss, p. 67 (non Helix deflexa Braun in Walchner, 1851). 894 1861 H.[elix] macrochila [sic] Reuss. – Reuss, p. 67. 1869a Helix (Crenea) expansilabris Sandberger. – Slavík, p. 245 (non Helix expansilabris Sandberger, 1858). 1869b Helix (Crenea) expansilabris Sandb. – Slavík, p. 271 (non Helix expansilabris Sandberger, 1858). 1870a Helix (Macularia Alb.) expansilabris Sandberger. – Boettger, p. 290 (non Helix expansilabris Sandberger, 1858). 1870a Helix (Macularia) bohemica; Boettger, p. 290, pl. 13, figs 4a–c. 1875 Helix bohemica Böttger. – Sandberger, p. 432, pl. 24, figs 8, 8a. 1891 Helix (Coryda) bohemica Boettger. – Klika, p. 57, text-figs 54a–c. 1891 Helix (Coryda) hortulana Thomae. – Klika, p. 59, text-figs 55a, b (non Helix hortulana Thomä, 1845). 1892 Helix (Coryda) bohemica Boettger. – Klika, p. 55, text-figs 54a–c. 1892 Helix (Coryda) hortulana Thomae. – Klika, p. 57, text-figs 55a, b (non Helix hortulana Thomä, 1845). 1897 Helix (Coryda) rugulosa var. subsulcosa Thom. – Babor, p. 17 (non Helix subsulcosa Thomä, 1845). 1909 [Tachea] bohemica O. Bttg. – Boettger, p. 18. 1911 [Helix] (Coryda, Macularia) bohemica Bttg. – Kafka, p. 68. 1911 Helix (Coryda) hortulana Th. – Kafka, p. 68 (non Helix hortulana Thomä, 1845). 1914 Cepaea hortulana (Tho.). – Wenz in Fischer & Wenz, p. 77 (pars). 1916 [Helix] (Coryda) bohemica Boettger. – Thuma, p. 83. 1916 [Helix] (Coryda) hortulana Thomae. – Thuma, p. 83 (non Helix hortulana Thomä, 1845). ? 1916 [Helix] (Obba) cfr hyperbolica Sandberger. – Thuma, p. 83 [non Helix (Obba) hyperbolica Sandberger, 1872]. 1917 Cepaea bohemica (Boettger). – Wenz, p. 60. 1923 Cepaea bohemica (Boettger). – Wenz, p. 609 (cum syn.). 1930 Holcotachea bohemica (Bttg.). – Pfeffer, p. 158. 1930 Holcotachea bohemica var. hortulanaeformis nov.; Pfeffer, p. 159. 1980 [Cepaea bohemica] hortulaniformis [sic] G. Pfeffer, 1929. – Richardson, p. 268. 1980 [Cepaea] bohemica (Böttger, 1870). – Richardson, p. 268. ? 2006 Cepaea bohemica (Boettger), 1870. – Kókay, p. 93, pl. 36, fig. 1. non 1911 Helix (Coryda?) bohemica Bttg. – Gaál, p. 59, pl. 3, fig. 5. Material. – One specimen (NHMW 2013/0572/0069), 169 specimens (NHMW 1890/0013/0402, 1909/0001/0046, 1909/0001/0047, 1923/409, 1926/0002/1347), 5 specimens Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake from Lipno (NHMW 1864/0012/0720), one specimen (MB.Ga.12215, Museum für Naturkunde, Berlin; Fig. 17Q, R). Dimensions. – Diameter: 19 mm, height: 13.5 mm (Fig. 17M–P); diameter: 17.5 mm, height: 13 mm (Fig. 17S, T). Description. – A very frequent, medium-sized, globular to ovoid shell with very dense and fine growth lines. Colour pattern usually well-preserved, consisting of three moderately broad, dark bands on the lower part of the last whorl and the base; a narrower fourth one is occasionally preserved or developed some millimetres below the upper suture. Some specimens develop a rather high and moderately gradate spire; very large shells tend to form somewhat flaring outer lips and a strongly oblique apertural plane. is too poorly preserved to allow a clear identification. A poorly preserved late Middle Miocene (Sarmatian) shell from Hungary described by Gaál (1911) represents clearly another species. Class Bivalvia Linnaeus, 1758 Superorder Heterodonta Neumayr, 1883 Order Venerida Gray, 1854 Superfamily Sphaerioidea Deshayes, 1855 Family Sphaeriidae Deshayes, 1855 Genus Sphaerium Scopoli, 1777 Type species. – Tellina cornea Linnaeus, 1758; by monotypy (Welter-Schultes 2012). Recent, Europe. Discussion. – Boettger (1870a) introduced Helix (Macularia) bohemica for the globular morphotypes, which have been erroneously identified so far with Helix rostrata and H. deflexa by Reuss in Reuss & Meyer (1849b) and Reuss (1861). The high-spired morphotypes were treated as distinct species by Boettger (1870a) (“expansilabris Sandberger”). Later, these were mistaken for the Oligocene Palaeotachea hortulana (Thomä, 1845) by Klika (1891). Pfeffer (1930) recognised the mistake and proposed the variety name hortulanaeformis for these shells, overlooking that Reuss in Reuss & Meyer (1849) had already introduced Helix macrocheila for these shells. Pfeffer (1930) had only 16 “typical” shells and a single “hortulanaeformis” shell at hand. Therefore, he was not able to evaluate the variability of this species. Based on the large number of specimens available to us, we agree with Wenz (1923) and Richardson (1980) that these high-spired shells are only morphotypes of a single species, which is usually referred to as Cepaea bohemica (Boettger, 1870). By treating all these taxa from Tuchořice as a single species, Helix macrocheila Reuss in Reuss & Meyer, 1849 gains priority over Helix bohemica Boettger, 1870. This species is the type species of Holcotachea Pfeffer, 1930 (subsequent designation by Nordsieck 1986), which is considered a synonym of Megalotachea by Nordsieck (1986) referring to the descriptions of Pfeffer (1930). It is unclear, which taxon from Korozluky was confused with the Eocene “Helix” hyperbolica by Thuma (1916). Obviously it is extremely unlikely that this species occurs in the Miocene Most Basin (Oppenheim 1916) and most probably he had a globular, high-spired Megalotachea macrocheila at hand. Sphaerium prominulum (Reuss in Reuss & Meyer, 1849) Figure 18A–L Occurrence. – Known from Korozluky, Pyšná, Tuchořice and Lipno; the shell from the Lower Miocene of the Somlóvásárhely drilling in Hungary described by Kókay (2006) Material. – 15 specimens (NHMW 1909/0001/0096, 1909/0001/0097), 3 specimens in the Prague collection (NM-PM-P 512–514). 1849a Cyclas prominula Rss.; Reuss, p. 12 (nomen nudum). 1849a Cyclas seminulum Rss.; Reuss, p. 12 (nomen nudum). 1849b Cyclas cornea L. (Lam.). – Reuss in Reuss & Meyer, p. 41, pl. 4, fig. 13 (non Tellina corneum Linnaeus, 1758). *1849a C.[yclas] prominula m.; Reuss, p. 42, pl. 4, fig. 14. 1849a C.[yclas] seminulum m.; Reuss, p. 42, pl. 4, fig. 15. 1861 Cyclas pseudocornea Reuss; Reuss, p. 82. 1861 C.[yclas] prominula Reuss. – Reuss, p. 82. 1861 C.[yclas] seminulum Reuss. – Reuss, p. 82. 1869a Sphaerium pseudocorneum Reuss. – Slavík, p. 268. 1869a Sphaerium prominulum Reuss. – Slavík, p. 268. 1869a Sphaerium seminulum Reuss. – Slavík, p. 268. 1869b Sphaerium pseudocorneum Rss. – Slavík, p. 272. 1869b Sphaerium prominulum Rss. – Slavík, p. 272. 1869b Sphaerium seminulum Rss. – Slavík, p. 272. 1870 Cyclas pseudocornea Rss. – Boettger, p. 299. 1870 Cyclas prominulum Rss. – Boettger, p. 299. 1870 Cyclas seminulum Rss. – Boettger, p. 299. 1875 Sphaerium pseudocorneum Reuss. – Sandberger, pp. 366, 423. 1891 Sphaerium pseudocorneum Reuss. – Klika, p. 115, text-figs 115a, b. 1892 Sphaerium pseudocorneum Reuss. – Klika, p. 110, text-figs 115a, b. 1916 Sphaerium pseudocorneum Reuss. – Thuma, p. 83. 895 Bulletin of Geosciences • Vol. 89, 4, 2014 A D B C E F G I H J K L Figure 18. Sphaerium prominulum (Reuss in Reuss & Meyer, 1849). • A – NM-PM-P 513 (bivalved adult specimen, left valve up). • B, G – NHMW 1909/0001/0097 (adult specimen, right valve). • C – NM-PM-P 512 (adult specimen, left valve). • D, J – NHMW 1909/0001/0096 (juvenile specimen, right valve). • E, H, K – NHMW 1909/0001/0096 (juvenile specimen, right valve). • F, I, L – NHMW 1909/0001/0096 (juvenile specimen, left valve). Dimensions. – The only adult specimen in the NHMW collection measures 7.5 mm in lenght, 6.5 mm in height and 2.1 mm in width (right valve, Fig. 18B, G). Description. – Oval, symmetrical shell with anterior part slightly larger and elongate than posterior portion. Hinge very thin and delicate in both valves. Right valve: C3 thin, arcuate, with thin anterior portion and thickened posterior portion, measuring near 200 μm, flanked by dorsal and ventral furrows (for C2 and C4); A1 short, thin, arcuate; P1 likewise thin but more elongated; A3 and P3 very small and thin, almost indiscernable even in SEM images. Left valve: C2 prominent, leaf-like, arcuate, ca 100 μm in 896 length; C4 elongate, with narrow anterior and thickened posterior portion, near 100 μm in length; both A2 and P2 prominent, short, triangular. Ligamental pit narrow, lanceolate, about 500 μm in length. Inner shell surface densely covered with small circular pores (ca 5 μm in diameter). Outer shell surface covered with dense, faint growth lines; occasionally growth irruglarities may result in local thickenings; no distinct transition between juvenile and adult shell. Discussion. – Reuss (1861) introduced Cyclas pseudocornea as new name for the specimen, which he erroneously had described as Cyclas cornea in 1849. At that time he still considered the two other species C. prominulum and Mathias Harzhauser et al. • The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake C. seminulum as distinct species. Later, Boettger (1970) and Sandberger (1875) considered the three taxa described by Reuss in Reuss & Meyer (1849b) from Tuchořice to be conspecific, representing different ontogenetic stages. We follow this view, which makes pseudocorneum a younger subjective synonym of prominulum and seminulum. As First Reviser we choose Sphaerium prominulum as name. Occurrence. – Known from Tuchořice, Jirkov, Korozluky and Lipno in the Most Basin. Additional occurrences from Hochheim, Gamerschwang, Öpfingen, Kaltennordheim, Theobaldshof/Rhön and Ulm, listed by Sandberger (1875), Klika (1891), and Fischer & Wenz (1915) need confirmation. Dubious taxa and species inquirendae The following taxa are either 1. based on insufficient material but might be valid species, 2. have been only mentioned in papers on the Most Basin fauna and might represent misidentifications, or 3. are based on unidentifiable fragments of (probably) other species. Material. – One fragment in the Prague collection (NM-PM-P 679). Dimensions. – Diameter: 2 mm. Discussion. – Only a fragment of the last two whorls was available to Klika (1891). The internal mould is attached to the matrix along the aperture. Therefore, any identification is impossible but the fragment may simply represent one of the Pseudoleacina species. The genus Opeas Albers, 1850, with its type species Helix goodalli Miller, 1822 (subsequent designation by Martens in Albers & Martens, 1860; Recent, Cuba), is known from the Early and Middle Miocene of Europe (Schlickum 1964). The only known European representative is Opeas minutum (Klein, 1853), which differs considerably in its much shorter last whorl and characteristic serrated suture (Harzhauser et al. 2014). We strongly doubt that the Bohemian specimen belongs to Opeas. Occurrence. – Only known from Pyšná. “Clausilia peregrina Reuss in Reus & Meyer, 1849” Oxychilus thomaeanum (Braun in Walchner, 1851) 1845 Helix deplanata Nob. – Thomä, p. 146 (non Helix deplanata Müller). *1851 Helix Thomaeana A. Braun; Braun in Walchner, p. 1140, p. 56 in offprint, No. 345. 1891 Hyalinia (Polita) deplanata Tho. – Boettger, p. 230 (non Helix deplanata Müller). 1917 Hyalinia (Hyalinia) thomaeana Wenz n. nom.; Wenz, p. 55. 1923 Oxychilus thomaeanum (A. Braun). – Wenz, p. 285 (cum syn.). Discussion. – This Early Miocene species was mentioned by Boettger (1891) only “en passant” from Tuchořice in his book-review of Klika (1891). Since then, no additional shell could be detected. This record might either represent a misidentification or an extremely rare species. Braun in Walchner (1851) recognised that Helix deplanata Thomä, 1845 was preoccupied and introduced the replacement name Helix Thomaeana. There is no reason to consider this name a nomen nudum as done by Wenz (1917) and the authorship has to be passed to Braun in Walchner (1851). “Opeas corruptum Klika, 1891” *1891 Opeas? corrupta n.; Klika, p. 71, text-fig. 67. 1911 Opeas? corrupta Kl. – Kafka, p. 68. 1823 ?Opeas corrupta Klika. – Wenz, p. 872 (cum syn.). 1849a Clausilia peregrina Rss.; Reuss in Reuss & Meyer, p. 11 (nomen nudum). *1849b C.[lausilia] peregrina m.; Reuss in Reuss & Meyer, p., 34, pl. 4, fig. 2. 1861 Cl.[ausilia] peregrina Reuss. – Reuss, p., 77. 1877 Cl.[ausilia] peregrina Reuss. – Boettger, p. 111. Discussion. – Is a nomen dubium (Boettger 1877). “Acrochasma tricarinatum Reuss, 1861” 1861 A.[crochasma] tricarinatum Reuss; Reuss, p. 80, pl. 3, fig. 16. 1891 Acrochasma tricarinatum Reuss. – Klika, p. 111, text-figs 110a–c. 1892 Acrochasma tricarinatum Reuss. – Klika, p. 106, text-figs 110a–c. Discussion. – This strange structure is clearly not a gastropod and is probably not even a Miocene fossil. Acknowledgements We are greatly indebted to Hartmut Nordsieck (Senckenberg Naturmuseum, Frankfurt). This paper benefitted enormously from his in-depth review and his expertise on Eurasian terrestrial gastropods. We thank Zlatko Kvaček (Charles University, Insti- 897 Bulletin of Geosciences • Vol. 89, 4, 2014 tute of Geology and Palaeontology, Prague) for help with Most Basin stratigraphy and Jiří Kvaček, Kamil Zágoršek and Eva Kadlecová (National Museum Prague) for sending rare literature and for providing access to the collections of the National Museum in Prague. Many thanks go to Anita Eschner and Sandra Kirchner (all NHM Vienna) for providing access to the malacological collections and pictures and to Martin Aberhan (Museum für Naturkunde, Berlin) for sending pictures of type material from the palaeontological collection in Berlin. Eike Neubert (Naturhistorisches Museum Bern) helped with critical comments on “Napaeus”. We are especially grateful to Alexander Nützel (Bayerische Staatssammlung für Paläontologie und Geologie, Munich), Daniela Esu (Università “La Sapienza” Roma), Ewa Stworzewicz (Polish Academy of Sciences, Kraków), Giuseppe Manganelli (Università di Siena) and Ronald Janssen (Senckenberg Naturmuseum, Frankfurt) for their careful reviews and constructive suggestions. Sigrid Hof (Sektion Malakologie, Senckenberg Naturmuseum, Frankfurt) provided pictures of Miophaedusa perforata. 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