Early Miocene Continental Gastropods from New Localities of the Molasse Basin in Southern Germany

PalZ (2016) 90:469–491 DOI 10.1007/s12542-016-0291-y

RESEARCH PAPER

Early Miocene continental gastropods from new localities of the Molasse Basin in southern Germany

1,2 3 3 Rodrigo B. Salvador • Martina Pippe`rr • Bettina Reichenbacher • Michael W. Rasser1

Received: 23 February 2015 / Accepted: 17 January 2016 / Published online: 12 February 2016 Ó Pala¨ontologische Gesellschaft 2016

Abstract Here, we present the continental aquatic and latter can be considered a synonym of T. cyrtocelis). terrestrial gastropods found in samples of 11 new boreholes Finally, we present a paleoecological interpretation for the in the Molasse Basin, southern Germany. The samples USM, OBM and OSM based on the gastropod fauna. come from the Lower Freshwater Molasse (USM), the Upper Brackish Molasse (OBM; Grimmelfingen and Keywords Gastropoda Á Grimmelfingen Formation Á Kirchberg Formations) and the Upper Freshwater Molasse Kirchberg Formation Á Lower Freshwater Molasse Á Upper (OSM). The studied segments of these lithostratigraphical Brackish Molasse Á Upper Freshwater Molasse units represent the beginning of the lower Miocene (USM), and the uppermost lower Miocene (OBM, OSM). Twenty- Kurzfassung Basierend auf den Proben von 11 in den four species of terrestrial and freshwater gastropods are letzten Jahren durchgefu¨hrten Bohrungen im su¨ddeutschen reported here, belonging to the families Neritidae, Molassebecken wird die kontinentale aquatische und ter- Melanopsidae, Pachychilidae, Bithyniidae, Hydrobiidae, restrische Gastropodenfauna neu bearbeitet. Das Material Truncatellidae(?), Viviparidae, Valvatidae(?), Lymnaeidae, stammt aus Proben der Unteren Su¨ßwassermolasse (USM), Planorbidae, Carychiidae, Zonitidae, Helicidae, Hygromi- der Oberen Brackwassermolasse (OBM; Grimmelfingen idae and Discidae(?). We provide remarks on the taxonomy und Kirchberg Formationen) und der Oberen Su¨ßwasser- of some of the studied species, including a revision of molasse (OSM). Die hier bearbeiteten Abschnitte dieser Theodoxus cyrtocelis, T. obstusangula and T. sparsus (the lithostratigraphischen Einheiten sind in das unterste Unter- Mioza¨n (USM) bzw. in das oberste Unter-Mioza¨n (OBM, OSM) einzustufen. Insgesamt werden 24 Arten kontinen- taler Gastropoden aus den Familien Neritidae, Melanopsi- dae, Pachychilidae, Bithyniidae, Hydrobiidae, Truncatellidae(?), Viviparidae, Valvatidae(?), Lymnaeidae, Planorbidae, Carychiidae, Zonitidae, Helicidae, Hygromi- Electronic supplementary material The online version of this idae und Discidae(?) beschrieben. Die Taxonomie einiger article (doi:10.1007/s12542-016-0291-y) contains supplementary material, which is available to authorized users. dieser Arten wird kritisch beleuchtet und die Arten der Gattung Theodoxus (T. cyrtocelis, T. obstusangula, T. & Rodrigo B. Salvador sparsus) warden revidiert, T. sparsus wird als Synonym [email protected] von T. cyrtocelis interpretiert. Daru¨ber hinaus wird auf der Grundlage der Gastropodenfauna eine palaöo¨kologische 1 Staatliches Museum fu¨r Naturkunde Stuttgart, Stuttgart, Germany Interpretation fu¨r die USM, OBM und OSM gegeben. 2 Mathematisch-Naturwissenschaftliche Fakulta¨t, Eberhard Karls Universita¨tTu¨bingen, Tu¨bingen, Germany Schlu¨sselwo¨rter Gastropoda Á Grimmelfingen Formation Á Kirchberg Formation Á Untere 3 Department of Earth and Environmental Sciences, Palaeontology and Geobiology, Ludwig-Maximilians- Su¨ßwassermolasse Á Obere Brackwassermolasse Á Obere Universita¨tMu¨nchen, Munich, Germany Su¨ßwassermolasse 123 470 R. B. Salvador et al.

Introduction R. Schlickum in the 1960s and 1970s, and some works dedicated to special aspects such as ontogenetic strategies Numerous samples from the cores of 11 boreholes located (Kowalke and Reichenbacher 2005), biogeographic distri- in the Molasse Basin of Bavaria, southern Germany bution (e.g., Reichenbacher 1993) or community studies (Fig. 1), have recently been investigated in the course of a (Schneider and Prieto 2011). The newly available material research program of the Bayerisches Landesamt fu¨r from the drilling research program mentioned above pro- Umwelt (project ‘‘Informationsoffensive Oberﬂa¨chennahe vides a unique opportunity to present a comprehensive Geothermie’’). The sampled lithostratigraphic units are taxonomical study of all the brackish, freshwater and ter- known as the Lower Freshwater Molasse (USM), the Upper restrial gastropods from the USM, OBM and OSM, and Brackish Molasse (OBM, Grimmelﬁngen and Kirchberg allows new paleoecological considerations. It was also Formations [Fms.]) and the Upper Freshwater Molasse possible to include type material for some species in order (OSM, Fig. 2). The studied samples from the USM are to achieve a better understanding of their taxonomy. dated from the lowermost Miocene (Aquitanian), while those from the OBM and the OSM correspond to the uppermost lower Miocene (upper Burdigalian). Geological setting Whereas the molluscan fauna from the USM has not yet been studied, several works are available for the OBM and The boreholes are situated in the western to central part of USM including the pioneering works of nineteenth century the Molasse Basin (Fig. 1), which is part of the North paleontologists (e.g., Krauss 1852), the later works of W. Alpine Foreland Basin and belongs to the Paratethys realm.

Fig. 1 Map showing the localities studied in the present work. a Schematic map of the Molasse Basin (modiﬁed after Jin et al. 1995). b Geographic locations of the boreholes

123 Early Miocene continental gastropods from new localities of the Molasse Basin in southern… 471

Fig. 2 Miocene lithostratigraphic units in the south German Molasse Basin (after Doppler et al. 2005; Reichenbacher et al. 2013) and stratigraphic range of drill cores. Dotted lines indicate stratigraphic units not present in the drill cores

In the German segment of the North Alpine Foreland et al. 1998). The thick marls of the Kirchberg Fm. (up to Basin, the Molasse succession as a whole is divided into 22 m) overlie the Grimmelfingen Fm. without a disconti- two transgressive–regressive megacycles, which are sepa- nuity. Typically, these marls contain rich and diverse fossil rated by an unconformity (e.g., Lemcke 1988; Bachmann assemblages with bivalves, gastropods, ostracods, fish oto- and Mu¨ller 1992; Schwerd et al. 1996; Doppler et al. 2005). liths and charophytes that allow for a subdivision into the The first megacycle, spanning the lower Oligocene (Ru- brackish–marine lower Kirchberg Fm. and the brackish– pelian) to lowermost Miocene (Aquitanian), comprises the lacustrine upper Kirchberg Fm. (Reichenbacher 1989; Lower Marine Molasse, the Lower Brackish Molasse, and Doppler 2011; Reichenbacher et al. 2013). the USM. The second megacycle includes the Upper Reichenbacher et al. (2013) have suggested an upper Marine Molasse (OMM), the OBM and OSM deposits and lower Miocene (upper Burdigalian) age for the OBM sed- is late Early Miocene (latest Aquitanian and Burdigalian) iments, based on a combination of bio-, litho- and magne- to early Late Miocene (Tortonian) in age. tostratigraphy, with the Kirchberg Fm. representing the This study focuses on the USM, OBM (Grimmelfingen uppermost Burdigalian (lower Karpatian). The OBM/OSM and Kirchberg Formations) and OSM gastropods (note that, transition is gradual in all drill cores. In most of the cores, here, we opt for the more widely used German abbrevia- only the lowermost OSM sediments, which are middle tions; USM: ‘‘Untere Su¨ßwassermolasse’’; OBM: ‘‘Obere Karpatian in age (Reichenbacher et al. 2013), contain few Brackwassermolasse’’; OSM: ‘‘Obere Su¨ßwassermolasse’’; freshwater fossils and rare brackish microfossils. Fig. 2). The USM sediments of the drill cores have yielded partially rich fossil assemblages with charophytes, gastropods, ostracods, fish otoliths and small mammal teeth, Materials and methods which are most probably late Egerian (earliest Miocene, Aquitanian) in age (unpublished data). Our new material has been assembled from the cores of The Grimmelfingen and Kirchberg Formations are gen- several boreholes drilled under the auspices of the Bavarian erally summarized as OBM. The Grimmelfingen Fm. over- Environment Agency (Landesamt fu¨r Umwelt) between lies discordantly the USM (Fig. 2) and is between 7 and 2010 and 2013 (Offingen 1, Glo¨tt 1, Gundremmingen 1, 41 m thick. This discordancy is due to the incision of a river Holzheim 1, Dillingen 1, Buttenwiesen 1, Druisheim 1, (Graupensand River) that extended along the northern Hamlar 1) as well as from the boreholes ‘‘Gu¨nzburg margin of the Southwest German Molasse Basin and eroded Brunnen 3 neu’’, ‘‘Vogelho¨lzl N Lauingen’’ and ‘‘WWA the entire OMM and partly also the upper USM in this area Donauwo¨rth, Kernbohrung Gempfing W4’’. (Kiderlen 1931; Reichenbacher et al. 1998). The Grim- Core samples were processed by soaking in hydrogen melfingen Fm. represents deposition of the Graupensand peroxide solution for several hours, washing through sieves River and consists of fine- to coarse-grained sands that are of 63-, 200- and 400-lm mesh size, and drying at a tem- predominantly carbonate- and fossil-free; only a few beds perature of 40 8C. Identification of USM, OBM and OSM contain mollusks, fish teeth and charophytes (Reichenbacher in borehole segments and samples has been conducted

123 472 R. B. Salvador et al. based on litho and biofacies as well as taxonomic and Family Neritidae Rafinesque, 1815 biostratigraphic analyses involving presence/absence of Genus Theodoxus Montfort, 1810 index taxa and characteristic fossil assemblages (including Theodoxus cyrtocelis (Krauss, 1852) charophytes, bivalves, gastropods, ostracods, fish otoliths Figure 3a–g. and mammal teeth). Details on the boreholes Hamlar 1, *1852 Neritina cyrtocelis Krauss: p. 145. Druisheim 1 and Gempfing W4 are available from *1852 Neritina sparsa Krauss: p. 145. Reichenbacher et al. (2013); publications dealing with the 1929 Theodoxus (Theodoxus) cyrtocelis cyrtocelis – Wenz: other boreholes are in preparation. The total numbers of p. 2991. samples and the number of gastropod-yielding samples that 1929 Theodoxus (Theodoxus) cyrtocelis sparsus – Wenz: were the basis for this study can be found in Online Sup- p. 2992. plementary Appendix 1. All cores are stored in the archives 1963 Theodoxus (Theodoxus) cyrtocelis cyrtocelis – Sch- of the Bavarian Environment Agency (Landesamt fu¨r lickum: p. 2. Umwelt) in Hof. 1964 Theodoxus (Theodoxus) cyrtocelis – Schlickum: p. 4, The fossil material is housed in the micropaleontology pl. 1, figs. 3–6. collection (Mikropalaöntologische Sammlung) of the 1966 Theodoxus (Theodoxus) cyrtocelis – Schlickum: p. 3, Bayerisches Landesamt fu¨r Umwelt in Munich, Germany, pl. 12, figs. 1–5. under the single record number LfU-SPR2014. The num- 1970a Theodoxus (Theodoxus) cyrtocelis – Schlickum: bers for the figured material received additional digits; the p. 146, pl. 10, figs. 3–4. remainder can be identified (species, locality, height in 1970b Theodoxus cyrtocelis – Schlickum: p. 162. sediment) via labels. Below, we indicate the occurrence of 1970c Theodoxus cyrtocelis – Schlickum: p. 176, pl. 3, each species regarding the borehole localities and main Fig. 1. stratigraphic units; for the detailed stratigraphical position 1989 Theodoxus cyrtocelis cyrtocelis – Reichenbacher: of each species in each site/borehole, please refer to the p. 144, pl. 1, figs. 1–2. tables in Online Supplementary Appendix 1. 1993 Theodoxus cyrtocelis cyrtocelis – Reichenbacher: Gastropod classification follows Bouchet et al. (2005), Tables 7, 13. with further modifications pointed out by Nordsieck (2014) 2005 Theodoxus cyrtocelis – Kowalke and Reichenbacher: for the European fossil land snails. All species found in the p. 617, figs. 4.3–4.7 material are figured and further data on their diagnostic 2014 Theodoxus (Theodoxus) cyrtocelis – Neubauer et al.: features is provided in the remarks section of each; full supplementary material 1. descriptions are provided only when required for further 2014 Theodoxus (Theodoxus) cyrtocelis sparsus – Neu- remarks on the species’ taxonomy. bauer et al.: supplementary material 1. The following abbreviations are used throughout the article. Institutional: LfU, Bayerisches Landesamt fu¨r Occurrence USM: Gundremmingen. OBM (Kirchberg Umwelt (Hof, Germany); MUWI, Naturwissenschaftliche Fm.): Buttenwiesen, Druisheim, Gempfing, Gu¨nzburg, Sammlung des Museums Wiesbaden (Wiesbaden, Ger- Hamlar 1, Holzheim, Lauingen. OSM: Glo¨tt (immediately many); NHMW, Naturhistorisches Museum Wien (Vienna, above the boundary with the Kirchberg Fm., so it could Austria); SMNS, Staatliches Museum fu¨r Naturkunde represent reworked material). Stuttgart (Stuttgart, Germany). Shell measurements: H, Description Shell with few quickly growing whorls. shell height; D, shell width (diameter); h, aperture/oper- Protoconch (*1 whorl) smooth, rounded, large (relative to culum length; d, aperture/operculum width. The specimens whole shell); transition to teleoconch clearly marked. were measured either with a digital caliper or with the aid Teleoconch sculptured by well-marked growth lines. Shell of computer software (Leica Application Suite [LAS] color from almost completely brown to almost completely v.3.8.0 and ImageJ). Stratigraphic units: USM, Lower white, with numerous patterns of white and brown dots and Freshwater Molasse; OBM, Upper Brackish Molasse; blotches. Suture shallow, but well-marked. Whorl profile OSM, Upper Freshwater Molasse. convex. Apex immersed to weakly raised. Peristome simple, sharp. Aperture D-shaped, with a large callus pad, which may show several very faint furrows (likely Systematic paleontology increasing with age). Operculum D-shaped, typical for genus, with an internal scimitar-shaped ridge. Phylum Mollusca, 1758 Class Gastropoda Cuvier, 1795 Remarks The present specimens compare very well to T. Neritimorpha Golikov and Starobogatov, 1975 cyrtocelis, a species originally described from the Kirch- Superfamily Neritoidea Rafinesque, 1815 berg Fm. (Krauss 1852). The type series (SMNS 123 Early Miocene continental gastropods from new localities of the Molasse Basin in southern… 473

Fig. 3 Aquatic gastropods. a, b Theodoxus cyrtocelis, in apertural D = 2.4 mm). h, i Theodoxus cyrtocelis, syntype, in apertural and and dorsal views, respectively (LfU-SPR2014-015; Hamlar 1, OBM, dorsal views, respectively (SMNS 25291/2005; Kirchberg an der Iller, Kirchberg Fm.; H = 6.7 mm, D = 7.4 mm). c, d Theodoxus cyrto- OBM, Kirchberg Fm.; D = 7.5 mm). j, k Theodoxus obtusangula,in celis, in apertural and dorsal views, respectively (LfU-SPR2014-016; apertural and dorsal views, respectively (LfU-SPR2014-014; Butten- Hamlar 1, OBM, Kirchberg Fm.; H = 6.4 mm, D = 6.6 mm). wiesen, OBM, Kirchberg Fm.; H = 5.0 mm, D = 4.9 mm). l, e Theodoxus cyrtocelis, dorsal to apical view (LfU-SPR2014-017 m Theodoxus obtusangula, syntype, in apertural and dorsal views, spcm. #1; Hamlar 1, OBM, Kirchberg Fm.; H = 5.0 mm, respectively (SMNS 25294/2005; Kirchberg an der Iller, OBM, D = 4.9 mm). f Theodoxus cyrtocelis, dorsal view (LfU-SPR2014- Kirchberg Fm.; D = 8.1 mm). n Theodoxus sparsus, syntype, in 017 spcm. #2; Hamlar 1, OBM, Kirchberg Fm.; H = 4.5 mm, dorsal to apical view (SMNS 25300/2005; Kirchberg an der Iller, D = 5.0 mm). g Theodoxus cyrtocelis, operculum (LfU-SPR2014- OBM, Kirchberg Fm.; D = 5.7 mm) 021; Buttenwiesen, OBM, Grimmelﬁngen Fm.; H = 4.6 mm,

25291/2005; 17 specimens; Fig. 3h, i) of this species show rounder or slenderer shells). The same variation of shape a good range of variation in color pattern (from completely (mainly regarding higher or lower spires, see below) and an brown shells to white and brown to almost completely even greater variation in color pattern can be seen in the white) and overall shell shape (e.g., higher or lower spires, present specimens. Theodoxus species are remarkable for

123 474 R. B. Salvador et al. exhibiting this kind of variation, as reported for many Fig. 4 Aquatic gastropods. a Melanopsis impressa, spire apex c recent representatives of the genus (e.g., Heller 1979; fragment, apertural view (LfU-SPR2014-007; Offingen, OBM, Kirch- berg Fm.; H = 6.0 mm). b Melanopsis impressa, aperture fragment Bandel 2001; Anistratenko 2005; Gloër and Pesˇic 2015). (LfU-SPR2014-008; Hamlar 1, OBM, Kirchberg Fm.; H = 10.9 mm). c Tinnyea escheri, apertural view (LfU-SPR2014- The opercula found (Fig. 3g) come from samples 004; Lauingen, OBM, Kirchberg Fm.; H = 9.7 mm). d Tinnyea including only T. cyrtocelis (Hamlar 1), so they can be escheri, apertural view (LfU-SPR2014-005; Hamlar 1, OBM, Kirch- reasonably assigned to this species. berg Fm.; H = 12.1 mm). e Bithynia glabra, apertural view (LfU- Krauss (1852) described another Theodoxus species SPR2014-020; Glo¨tt, OBM, Kirchberg Fm.; H = 9.1 mm). f Bithynia glabra, apertural view (LfU-SPR2014-020; Glo¨tt, OBM, Kirchberg from the Kirchberg Fm., namely T. sparsus, which he Fm.; H = 8.1 mm). g Bithynia glabra, apertural view (LfU- defined by a flattened spire and gray color (syntypes: SPR2014-019 spcm. #1; Buttenwiesen, OBM, Kirchberg Fm.; SMNS 253000/2005, 7 specimens; Fig. 3n). We consider H = 8.9 mm). h Bithynia glabra, specimen more akin to B. dunkeri, here T. sparsus a synonym of T. cyrtocelis for the following apertural view (LfU-SPR2014-019 spcm. #2; Buttenwiesen, OBM, Kirchberg Fm.; H = 9.4 mm). i Bithynia glabra, operculum (LfU- reasons. The gray color seen in Krauss’ shells is the same SPR2014-040; Druisheim, OBM, Kirchberg Fm.; H = 4.2 mm, as the rock matrix and is probably a preservation artifact; D = 3.5 mm). j Ctyrokya conoidea, apertural view (LfU-SPR2014- one of the syntypes actually is white and brown, like typ- 037; Hamlar 1, OBM, Kirchberg Fm.; H = 4.1 mm). k Cty- ical T. cyrtocelis. The flattened spires can be considered as rokya conoidea, with preserved color pattern, apertural view (LfU- SPR2014-013; Hamlar 1, OBM, Kirchberg Fm.; H = 3.5 mm). variation in overall shell shape, as known from recent l Hydrobia semiconvexa, apertural view (LfU-SPR2014-031; Hamlar Theodoxus species (e.g., Elkarmi and Ismail 2005; Gloër 1, OBM, Kirchberg Fm.; H = 5.0 mm). m Nematurella bavarica, and Pesˇic 2015). Moreover, some syntypes of T. sparsus apertural view (LfU-SPR2014-047 spcm.. #1; Gu¨nzburg, OBM, have more pronounced spires while some syntypes of T. Kirchberg Fm.; H = 3.9 mm). n Nematurella bavarica, apertural view (LfU-SPR2014-042 spcm.. #1; Druisheim, OBM, Grimmelfin- cyrtocelis (SMNS 25291/2005) have flattened spires. gen Fm.; H = 3.6 mm). o Nematurella bavarica, apertural view Finally, the overall impression of a more flattened spire (LfU-SPR2014-029; Lauingen, OBM, Kirchberg Fm.; H = 4.2 mm). profile can be the result of all specimens of T. sparsus p Nematurella bavarica, apertural view (LfU-SPR2014-042 spcm.. being either juvenile or fragmentary shells. As seen in the #2; Druisheim, OBM, Grimmelfingen Fm.; H = 3.6 mm). q Nema- turella bavarica, apertural view (LfU-SPR2014-043; Hamlar 1, recent T. macri (Sowerby, 1849), for instance, the overall OBM, Kirchberg Fm.; H = 4.3 mm). r Nematurella bavarica, shell shape and its proportions depend on age: the shells apertural view (LfU-SPR2014-047 spcm.. #2; Gu¨nzburg, OBM, become taller, changing the impression of a flattened spire Kirchberg Fm.; H = 4.2 mm). s Nematurella bavarica, in apertural to to a more prominent one, as the animals grow older and lateral view, showing circular hole, likely a predation mark (LfU- SPR2014-041; Buttenwiesen, OBM, Kirchberg Fm.; H = 3.5 mm). larger and, thus, add more whorls (Elkarmi and Ismail t ?Truncatella sp., apertural view (LfU-SPR2014-009; Hamlar 1, 2005). OBM, Kirchberg Fm.; H = 4.2 mm). u, v Viviparus cf. suevicus,in apertural and apical views, respectively (LfU-SPR2014-002, spcm. Theodoxus obtusangula (Krauss, 1852). #1; Druisheim, OBM, Grimmelfingen Fm.; H = 8.6 mm, Figure 3j, k. D = 9.9 mm). w ?Valvata sp., apertural view (LfU-SPR2014-001; *1852 Neritina obtusangula Krauss: p. 145. Druisheim, OBM, Grimmelfingen Fm.; H = 7.0 mm, D = 8.7 mm) 1929 Theodoxus (Theodoxus) cyrtocelis obtusangulus [sic] – Wenz: p. 2992. 2014 Theodoxus (Theodoxus) cyrtocelis obtusangulus [sic] Fm. (holotype: SMNS 106414; Fig. 3l, m). When com- – Neubauer et al.: supplementary material 1. pared to T. cyrtocelis, this species has a smaller proto- Occurrence OBM (Kirchberg Fm.): Buttenwiesen, conch, a smaller width of the initial whorls (which are also Gu¨nzburg. more tightly coiled and more regularly growing, but expanding suddenly around 2 whorls), a slightly larger Description Shell small, neritiform, with few regularly number of whorls, a well-defined angulation on upper growing whorls (but with a sudden expansion after *2 portion of whorl (which gives the species its name), a whorls). Protoconch (*1 whorl) smooth, relatively smaller usually uniformly brown color (Krauss had only one than in congeners; transition to teleoconch clear. Teleoconch specimen, but this seems to hold true for the present sculptured by well-marked growth lines. Shell color uni- specimens with a single exception; see description above) formly brown (a single specimen displays round white dots, and stronger and more well-marked growth lines. As such, similar to T. cyrtocelis). Suture shallow, but well-marked. with the presently available material, T. obtusangula seems Whorl profile slightly flattened, with well-defined angulation to be a clearly distinct species. on upper portion of whorl. Apex raised. Peristome simple, sharp. Aperture D-shaped, with large columellar region. The species epithet is derived from the Latin word ‘‘angulus’’ (=an angle). As such, ‘‘angula’’ is still a sub- Remarks Theodoxus obtusangula is the third Theodoxus stantive with a variant gender ending and is, therefore, not species described by Krauss (1852) from the Kirchberg declensable.

123 Early Miocene continental gastropods from new localities of the Molasse Basin in southern… 475

Caenogastropoda Cox, 1966 Melanopsis impressa Krauss, 1852 Superfamily Cerithioidea Fleming, 1822 Figure 4a, b. Family Melanopsidae H. Adams and Adams, 1854 *1852 Melanopsis impressa Krauss: p. 143, pl. 3, ﬁg. 3. Genus Melanopsis Fe´russac, 1807 1872 Melanopsis impressa – Sandberger: pl. 31, ﬁg. 8.

123 476 R. B. Salvador et al.

1872 Melanopsis impressa – Sandberger: p. 558. 1976 Brotia (Tinnyea) escheri – Schlickum: p. 4, pl. 1, 1964 Melanopsis impressa impressa – Schlickum: p. 11, pl. fig. 9. 2, figs. 23–25. 2000 Brotia (Tinnyea) escheri – Mikuzˇ and Pavsˇicˇ: p. 44, 1966 Melanopsis impressa impressa – Schlickum: p. 325, pl. 1, figs. 1–8. pl. 13, fig. 25. Occurrence OBM (Kirchberg Fm.): Buttenwiesen, 1970a Melanopsis impressa impressa – Schlickum: p. 148, Druisheim, Gu¨nzburg, Hamlar 1, Lauingen. pl. 10, fig. 5. 1970b Melanopsis impressa impressa – Schlickum: p. 160. Remarks Despite the very fragmentary nature of the 1973 Melanopsis impressa impressa – Steininger et al.: present material, the species’ turriform shell and charac- p. 404, pl. 4, figs. 6–7. teristic teleoconch sculpture (very strong opisthocline and 1989 Melanopsis impressa impressa – Reichenbacher: slightly curved axial ribs on first whorls and both axial ribs p. 174, pl. 2, fig. 6. and spiral cords on the remaining whorls) allow the iden- 1993 Melanopsis impressa impressa – Reichenbacher: tification. According to Harzhauser et al. (2002), the spe- Tables 6, 13. cies has a broad range of morphological plasticity, as 2001 Melanopsis impressa – Harzhauser and Kowalke: usually seen in recent Pachychilidae. p. 360, figs. 3.6–3.8. Tinnyea lauraea is known in the fossil record of Central 2005 Melanopsis impressa impressa – Kowalke and Europe from the Oligocene to the Pliocene (Harzhauser Reichenbacher: p. 830, figs. 8.3–8.5. et al. 2002). This large time span suggests that this is a still 2014 Melanopsis impressa – Neubauer et al.: supplemen- unresolved species complex (Kadolsky 1995) tary material 1. Superfamily Truncatelloidea Gray, 1840 Occurrence OBM (Grimmelfingen Fm.): Gu¨nzburg. Family Bithyniidae Gray, 1857 OBM (Kirchberg Fm.): Buttenwiesen, Druisheim, Glo¨tt, Genus Bithynia Leach, 1818 Gu¨nzburg, Hamlar 1, Holzheim, Offingen. Bithynia glabra (von Zieten, 1830) Remarks Theshellshaveaveryflatwhorlprofile,a Figure 4e, i slightly incised suture, a simple and sharp peristome and *1830 Cyclostoma glabrum von Zieten: p. 42, pl. 31, fig. 9. an elongated aperture, with a callus on the columellar 1852 Paludina tentaculata – Krauss: p. 140. and palatal regions; furthermore, the base of the aper- *1872 Bythynia [sic] gracilis – Sandberger: pl. 28, ture’s columellar region is folded, encircling the central figs. 16–16a. coiling axis of the shell and slightly bent inwards. These 1875 Bythynia [sic] gracilis – Sandberger: p. 561. features (especially the characteristic aperture, present 1964 Bithynia glabra – Schlickum: p. 9. even in juvenile specimens, such as the syntypes SMNS 1966 Bithynia dunkeri – Schlickum: p. 324, pl. 12, 106400) makes the species easily identifiable in the figs. 19–21. material. As already reported by Kowalke and 1966 Bithynia glabra – Schlickum: p. 324, pl. 12, fig. 22. Reichenbacher (2005), the width of the spire in this 1989 Bithynia cf. glabra – Reichenbacher: p. 144, pl. 1, species is slightly variable: shells with slender spires are fig. 4. more common, but some shells show wider and bulkier 1989 Bithynia dunkeri – Reichenbacher: p. 144, pl. 1, spires. figs. 6–7. 1993 Bithynia dunkeri – Reichenbacher: Tables 6–7. Melanopsis impressa is common in estuarine environments 1993 Bithynia glabra – Reichenbacher: Table 13. from the Miocene of the Paratethys and Mediterranean 2005 Bithynia cf. glabra – Kowalke & Reichenbacher: (Harzhauser and Kowalke 2001); it was originally descri- p. 629, figs. 8.1–8.2. bed from the Kirchberg Fm. of Kirchberg an der Iller 2014 Bithynia glabra – Neubauer et al.: supplementary (Krauss 1852). material 1. Family Pachychilidae P. Fischer and Crosse, 18 Occurrence USM: Offingen. OBM (Grimmelfingen Fm.): Genus Tinnyea Hantken, 1887 Buttenwiesen, Dillingen, Gu¨nzburg. OBM (Kirchberg Tinnyea lauraea (Mathe´ron, 1843) Fm.): Buttenwiesen, Dillingen, Druisheim, Gempfing, Figure 4c, d Glo¨tt, Gu¨nzburg, Hamlar 1, Holzheim, Lauingen, Offingen. OSM: Buttenwiesen, Dillingen, Gempfing, Glo¨tt, Gun- *1843 Melania lauraea Mathe´ron: p. 291, pl. 36, dremmingen, Hamlar 1, Holzheim, Offingen. figs. 23–24. 1927 Melanatria escheri turrita – Berz and Jooß: p. 206. Description Shell small, conical, imperforate. Protoconch 1953 Brotia (Tinnyea) escheri – Papp: p. 128, pl. 3, fig. 29. smooth (*1 whorl), rounded; transition to teleoconch not

123 Early Miocene continental gastropods from new localities of the Molasse Basin in southern… 477 clearly marked. Teleoconch smooth, except for growth 1965 Ctyrokya conoidea – Schlickum: p. 101. lines. Whorl profile strongly convex. Suture deep. Aperture 1973 Ctyrokya conoidea – Steininger et al.: p. 397, pl. 3, oval, pointed on the meeting of palatal and parietal regions. fig. 5. Peristome thickened, complete. Operculum with concentric 1989 Ctyrokya conoidea – Reichenbacher: p. 144, pl. 2, growth pattern and central nucleus. fig. 4. 2005 Ctyrokia conoidea [sic] – Kowalke and Reichen- Remarks Two species of Bithynia have been described bacher: p. 621, figs. 6.8–6.10. for the OBM: B. dunkeri Gude, 1913, from the Kirchberg 2014 Ctyrokya conoidea – Neubauer et al.: supplementary Fm. (but also occurring in the OSM), and B. glabra (von material 1. Zieten, 1830), from the Grimmelfingen and Kirchberg Fms. Schlickum (1966) notes that B. glabra substitutes B. dun- Occurrence OBM (Kirchberg Fm.): Buttenwiesen, keri in the younger layers of the Kirchberg Fm. at Druisheim, Gu¨nzburg, Hamlar 1, Holzheim, Lauingen, Leipheim. Offingen. OSM: Gundremmingen (at the lowermost layer, could be reworked material from older layers). There is some confusion in the literature about the identification of these two species. When only the early Remarks The shell is conical-elongated and broad, with a whorls are present, it is impossible to distinguish between flat whorl profile, a proportionately large body whorl, a them, while the adults differ according to the original large ellipsoid aperture, an incomplete peristome and a descriptions and later revision by Sandberger (1875): dome-shaped and prominent smooth protoconch (of ca. 1 Bithynia dunkeri has a broader shell, more flattened whorls, whorl) transitioning abruptly to a smooth teleoconch (ex- a proportionately larger body whorl (and thus proportion- cept for growth lines). The color pattern was apparently ately smaller spire) and a larger and more angular aperture; preserved in some specimens (Fig. 4k) and consists of Bithynia glabra is identified by its slenderer shell, pro- numerous intermittent dark spiral stripes. portionately smaller body whorl and very convex whorls. The broad conical spire of the present specimens, with flat However, this distinction might not be as clear-cut as whorls and a large aperture, compare well to the syntypes of previously supposed. C. conoidea (SMNS 106373), a species described by Krauss In the present material, there seems to be a gradation of (1852) from the Kirchberg Fm. Schlickum (1965) erected the forms from a more slender shell (i.e., more akin to B. genus Ctyrokya and included this species, but the validity of glabra; Fig. 4e) to a broader one (i.e., more akin to B. this genus remains a matter for future revisionary work. As dunkeri; Fig. 4h), with intermediate forms between them pointed out by Salvador et al. (in press), the substitution of (Fig. 4f, g). As such, the two described species could be the junior primary homonym is not mandatory if the condi- simply extreme forms in a continuum; moreover, the pro- tions of Article 23.9.5 ICZN are met. toconch size and its overall shape seem to be the same Two further species of Ctyrokya have been reported throughout this continuum in the present material. There is from the Kirchberg Fm., namely C. hoelzli (Schlickum, not a clear case of substitution of one species for the other 1964) and C. zoebeleini Schlickum, 1970a, b, c, d through the sedimentary layers; both forms are equally (Kowalke and Reichenbacher 2005), but they were not found throughout the Kirchberg Fm. (the only strati- found in the present material. graphical unit for which we have completely preserved specimens). Nevertheless, the present material is too scarce Genus Hydrobia Hartmann, 1821 and poorly preserved to delve deeper into this matter. A Hydrobia semiconvexa Sandberger, 1875 throrough taxonomical revision of these two species is Figure 4l urgently needed, with examination of the available type 1852 Litorinella acuta – Krauss: p. 142 [non Cyclostoma material and a large sample of topotypes. Meanwhile, we acutum Draparnaud, 1805]. prefer here the more cautious approach of a classification *1875 Hydrobia semiconvexa Sandberger: p. 561. as B. glabra, the older name. 1971 Hydrobia semiconvexa – Schlickum: p. 571, pl. 1, Family Hydrobiidae Stimpson, 1865 figs. 2–3. Genus Ctyrokya Schlickum, 1965 1973 Hydrobia semiconvexa – Steininger et al.: p. 391, pl. Ctyrokya conoidea (Krauss, 1852) 2, fig. 2. Figure 4j, k. 1989 Hydrobia semiconvexa – Reichenbacher: pl. 2, fig. 8. *1852 Paludina conoidea Krauss: p. 141, pl. 3, fig. 1 [non 1993 Hydrobia semiconvexa – Reichenbacher: p. 290. Paludina conoidea de Reynie`s, 1844:4;Ku¨ster, 1852: 43, 2005 Hydrobia cf. semiconvexa – Kowalke and Reichen- pl. 9, figs. 3–7]. bacher: p. 619, figs. 5.3–5.4.

123 478 R. B. Salvador et al.

Occurrence OBM (Kirchberg Fm.): Buttenwiesen, Fig. 5 Aquatic and terrestrial gastropods. a ?Valvata sp., apical view c Gempfing, Gu¨nzburg, Hamlar 1, Holzheim, Lauingen. (LfU-SPR2014-001; Druisheim, OBM, Grimmelfingen Fm.; H = 7.0 mm, D = 8.7 mm). b Stagnicola armaniacensis, apertural Remarks The narrow and conical-elongated shell, with a view (LfU-SPR2014-028; Offingen, OBM, Kirchberg Fm.; H = 3.9 mm). c Stagnicola praebouilleti, apertural view (LfU- prominent smooth protoconch (of ca. 1 whorl), a small SPR2014-026; Buttenwiesen, OBM, Grimmelfingen Fm.; ellipsoid aperture and a complete peristome, compares well H = 2.1 mm). d Lymnaea dilatata, apertural view (LfU-SPR2014- with the type specimens of H. semiconvexa (housed at the 018; Buttenwiesen, OBM, Kirchberg Fm.; H = 6.1 mm). e Ferris- MUWI). This species was originally described from the sia cf. wittmanni, apical view (LfU-SPR2014-025; Offingen, OBM, Kirchberg Fm.; H = 4.0 mm, D = 2.6 mm). f Gyraulus albertanus, Kirchberg Fm. from Illerkirchberg (Krauss 1852; Sand- apertural view (LfU-SPR2014-034 spcm. #1; Offingen, OBM, berger 1875). Kirchberg Fm.; H = 1.1 mm, D = 2.3 mm). g Gyraulus albertanus, umbilical view (LfU-SPR2014-034 spcm. #2; Offingen, OBM, However, the present specimens also compare reason- Kirchberg Fm.; D = 2.0 mm). h Gyraulus applanatus, apertural ably well to the syntypes H. frauenfeldi (Ho¨rnes, 1856) view (LfU-SPR2014-035; Buttenwiesen, OBM, Kirchberg Fm.; (NHMW 1846/0037/0456), but have a somewhat more H = 0.7 mm, D = 2.2 mm). i Gyraulus applanatus, umbilical view elongated aperture. This species was originally described (LfU-SPR2014-036; Offingen, OBM, Kirchberg Fm.; D = 2.4 mm). j Planorbarius mantelli, apical view (LfU-SPR2014-039; Offingen, from Sarmatian sediments at Hauskirchen, Austria (Ho¨rnes OBM, Kirchberg Fm.; D = 2.6 mm). k Carychium cf. galli, apertural 1856; Harzhauser and Piller 2004), but is also known from view (LfU-SPR2014-022; Buttenwiesen, OBM, Grimmelfingen Fm.; the OBM of the Oncophora Basin, which neighbors the H = 1.7 mm). l Archaeozonites sp., apical view (LfU-SPR2014-006; Kirchberg Basin (Schlickum 1964; Reichenbacher 1993; Offingen, OBM, Kirchberg Fm.; D = 6.1 mm). m–o Megalotachea cf. silvana, in apertural, apical and umbilical view, respectively (LfU- Kowalke and Reichenbacher 2005). A possible synonymy SPR2014-011; Glo¨tt, OSM; H = 14.9 mm, D = 23.4 mm). p–r between these two species should be further investigated. Palaeotachea sp., in apertural, apical and umbilical view, respectively (LfU-SPR2014-010; Dillingen, USM; H = 8.6 mm, D = 14.8 mm). Genus Nematurella Sandberger, 1875 s Leucochroopsis sp., apical view (LfU-SPR2014-024; Buttenwiesen, Nematurella bavarica Sandberger, 1875 OBM, Grimmelfingen Fm.; D = 4.3 mm). t, u ?Discus sp., in Figure 4m–s. apertural and apical view, respectively (LfU-SPR2014-023; Gun- *1875 Hydrobia bavarica Sandberger: p. 576. dremmingen, USM; H = 0.9 mm, D = 1.5 mm) *1960 Nematurella schuetti Schlickum: p. 211, pl. 19, fig. 12. 1961 Nematurella bavarica – Schlickum: p. 57. Occurrence OBM (Grimmelfingen Fm.): Buttenwiesen, 1963 Nematurella scholli – Schlickum: p. 3. Gu¨nzburg. OBM (Kirchberg Fm.): Buttenwiesen, Dillin- 1966 Nematurella zilchi – Schlickum: p. 223, pl. 12, gen, Druisheim, Gempfing, Glo¨tt, Gu¨nzburg, Hamlar 1, figs. 9–18. Holzheim, Lauingen, Offingen. OSM: Buttenwiesen, 1971 Nematurella convexula – Schlickum: pl. 1, fig. 5. Gempfing, Glo¨tt, Gundremmingen. 1973 Nematurella scholli – Steininger et al.: p; 391, pl. 2, Description Shell small, conical-elongated. Protoconch Fig. 5. (*1 whorl) prominent, rounded, dome-shaped, smooth; 1973 Nematurella convexula – Steininger et al.: p. 392, pl. transition to teleoconch clearly marked. Teleoconch 2, fig. 8. smooth, except for growth lines. Whorl profile convex. 1973 Nematurella zilchi – Steininger et al.: p. 393, pl. 2, Suture deep. Aperture ellipsoid. Peristome complete, fig. 10. slightly thickened. Umbilicus narrow, shallow. 1989 Nematurella convexula – Reichenbacher: p. 144, pl. 1, fig. 2. Remarks Several species of Nematurella have been 1989 Nematurella zilchi – Reichenbacher: p. 144, pl. 2, fig. 1. described from the OBM of the Kirchberg Basin and the 1989 Nematurella cf. bavarica – Reichenbacher: p. 144, pl. neighboring Oncophora Basin (e.g., Schlickum 1960; 2, fig. 3. Schlickum and Strauch 1967; Kowalke and Reichenbacher 2005 Nematurella bavarica – Kowalke and Reichenbacher: 2005). The first one to be described was N. bavarica, from p. 623, figs. 7.3–7.5. the lowermost OSM of Gu¨nzburg, with no figure and just a 2005 Nematurella convexula – Kowalke and Reichen- very brief description (Sandberger 1875). Analyzing what bacher: p. 627, figs. 5.9–5.10. is possibly original material from Sandberger, Schlickum 2014 Nematurella bavarica – Neubauer et al.: supple- (1961) synonymized one of his species, N. schuetti, with N. mentary material 1. bavarica. The figured type material of these many species 2014 Nematurella convexula – Neubauer et al.: supple- seem to be often indistinguishable from one another (e.g., mentary material 1. Schlickum 1960) and are usually what would be perceived 2014 Nematurella zilchi – Neubauer et al.: supplementary as a continuum of variation of a single species. Hydrobiids material 1. are remarkable in intraspecific conchological variability,

123 Early Miocene continental gastropods from new localities of the Molasse Basin in southern… 479

both hereditary and ecophenotypic (e.g., Dillon 2000; 3.6–4.3 mm (the protoconch of the Nematurella ‘‘species’’ Haase 2003; Vuolo et al. 2011). are also within a narrow and overlapping range; see Kowalke and Reichenbacher 2005) and range from slender This continuum of variation in shell shape is exactly to broad spires, from 4 to 5 whorls, from a deep suture what is seen in the present material (Fig. 4m–r): the (and, thus, more convex whorls) to a shallower suture (and specimens are all within a reasonable shell height range of

123 480 R. B. Salvador et al. more flattened whorls), etc. All these extremes, of course, Remarks The single, poorly preserved, juvenile specimen are accompanied by a series of intermediate forms in all the can be distinguished from all the other species described stratigraphical units where they occur. For example, the herein by its large and dome-shaped protoconch and its specimens of Fig. 4n would represent what is commonly pattern of growth, with the first whorls regularly growing identified as N. zilchi Schlickum, 1960 in the literature, and slowly increasing in width and an expanded body while those of Fig. 4q, r would be N. convexula Schlickum whorl. This specimen is very reminiscent of the genus and Strauch 1967; finally, those of Fig. 4o, p are clear Truncatella. However, its spire is slightly more conical intermediates that could be identified as either N. zilchi or than usual for the genus, though this could be due to the N. convexula. The broader shell of Fig. 4m is what corre- specimen being sub-adult. A more precise identification is sponds to typical N. bavarica. A future (and sorely needed) not possible at present. Few fossil Truncatella species are revision of this species complex should take into account known from western and central Europe, but the genus has this variability and intermediate forms, besides conducting records from the Paleocene to the Pliocene (Wenz 1928; a thorough analysis of the type material. Glibert 1973; Lozouet et al. 2001). Moreover, shell growth after breakage or abnormal Superfamily Viviparoidea Gray, 1847 growth on the body whorl are rather commonly found in Family Viviparidae Gray, 1847 the material and are responsible for alterations in overall Genus Viviparus Montfort, 1810 shell shape and proportions (for instance, fewer whorls, a Viviparus cf. suevicus Wenz, 1919 more truncated body whorl or a slightly detached aperture); Figure 4u, v these changed features seem to have been sometimes used for the diagnosis of new species of Nematurella (e.g., Occurrence OBM (Kirchberg Fm.): Druisheim. Schlickum 1960). Remarks Despite the poor preservation, the present Trace fossils Some specimens of N. bavarica (e.g., specimens can be assigned to the genus Viviparus based on Fig. 4s) and a single specimen of Stagnicola armaniacensis their size, the very characteristic shape of the first whorls (Noulet, 1857), described further below, show a circular and teleoconch sculpture pattern. Viviparus suevicus is the hole on their body whorls (a single specimen of N. only species of the genus recorded so far from the Kirch- bavarica shows two neighboring holes). These holes are berg Fm. (which is also its type stratum; type locality: the same as those reported by Rasser and Covich (2014) for Kirchberg an der Iller; Krauss 1852; holotype: SMNS the freshwater snails from Steinheim (Middle Miocene, 106437). Tinca SW Germany), related to predation by Cuvier, 1817 Heterobranchia Gray 1840 Tinca (Actinopterygii: Cyprinidae). The teeth of sp. Superfamily Valvatoidea Gray 1840 described from the Kirchberg Fm. (Reichenbacher 1988: Family Valvatidae Gray 1840 pl. 1, figs. 10–11) actually belong to another cyprinid, Genus Valvata Mu¨ller, 1773 ´ ¨ Palaeocarassius Obrhelova, 1970 (see Bohme 2010: ?Valvata sp Fig. 5a–c), but others cyprinids present in the Kirchberg Figures 4w, 5a Fm. (Reichenbacher 1988; Reichenbacher et al. 2004; Schulz-Mirbach and Reichenbacher 2006) could be Occurrence OBM (Kirchberg Fm.): Druisheim, Offingen. responsible for these marks. These ichnofossils are studied Remarks The overall shell shape (depressed trochiform, in more detail by Rasser et al. (in press). with round whorls rapidly growing in size, circular aperture Family Truncatellidae Gray 1840 and narrow umbilicus) and protoconch size and rounded shape Genus Truncatella Risso, 1826 seem consistent with the genus Valvata. The poor preservation ?Truncatella sp. prevents the observation of the protoconch sculpture, which is Figure 4t. very characteristic for the genus (Binder 1967;Riedel1993), precluding a more definitive generic assignment. Occurrence OBM (Kirchberg Fm.): Hamlar 1. Pulmonata Cuvier, 1814. Description Shell conical-elongated, with 4 whorls; Basommatophora Keferstein in Bronn, 1864 imperforate; regularly and slowly growing first whorls; Superfamily Lymnaeoidea Rafinesque, 1815 body whorl expanded. Protoconch large, rounded, dome- Family Lymnaeidae Rafinesque, 1815 shaped. Shell sculpture cannot be observed due to poor Genus Stagnicola Jeffreys, 1830 preservation. Whorls profile slightly convex. Suture well- Stagnicola armaniacensis (Noulet, 1857) marked but not incised. Aperture oval, elongated diago- Figure 5b. nally. Peristome complete.

123 Early Miocene continental gastropods from new localities of the Molasse Basin in southern… 481

*1852 Limnaea armaniacensis Noulet: p. 22. *1854 Limnea dilatata Noulet: p. 107. 1923 Galba (Galba) armaniacensis armaniacensis – 2000 Lymnaea dilatata – Fischer: p. 136, figs. 1–2. Wenz: p. 1351. 2002 Lymnaea dilatata – Binder: p. 165, pl. 1, fig. 7a. 1964 Stagnicola (Stagnicola) armaniacensis – Schlickum: 2004 Lymnaea dilatata – Binder: p. 192, pl. 1, fig. 7. p. 13, pl. 2, fig. 31. 2005 Lymnaea dilatata – Kowalke and Reichenbacher: 1989 Stagnicola armaniacensis – Reichenbacher: p. 174, p. 630, figs. 9.4–9.5. pl. 2, fig. 7. 2014 Lymnaea dilatata – Salvador and Rasser: p. 189, 1993 Stagnicola armaniacensis – Reichenbacher: figs. 8–9. Tables 7, 11–13. 2014 Radix dilatata – Neubauer et al.: supplementary 2005 Stagnicola armaniacensis – Kowalke and Reichen- material 1. bacher: p. 631, figs. 9.6–9.7. Occurrence USM: Gundremmingen, Offingen. OBM 2014 Stagnicola armaniacensis – Harzhauser et al.: 11, pl. (Grimmelfingen Fm.): Buttenwiesen. OBM (Kirchberg 2, figs. 4–11. Fm.): Buttenwiesen, Glo¨tt, Hamlar 1, Holzheim, Lauingen, 2014 Stagnicola armaniacensis – Neubauer et al.: supple- Offingen. OSM: Gempfing, Offingen. mentary material 1. Remarks Although only shell apexes remain, the spec- Occurrence USM: Gundremmingen, Offingen. OBM imens can be confidently assigned to Lymnaea dilatata. (Grimmelfingen Fm.): Buttenwiesen, Gu¨nzburg, Offingen. This species is diagnosed by its large size, its overall OBM (Kirchberg Fm.): Buttenwiesen, Dillingen, Glo¨tt, acuminated (although rather blunt on its very top) and Gu¨nzburg, Holzheim, Lauingen, Offingen. OSM: Gempf- proportionately short spire and the rapidly growing ing, Gundremmingen, Holzheim, Offingen. whorls. Originally described from Sansan (MN 6, Remarks The small and wide lymnaeid shells, with few France; see Fischer 2000), L. dilatata is known from the quickly growing whorls, a deep suture and an elongated oval whole Middle Miocene of West and Central Europe aperture, can be identified as S. armaniacensis. This species (Binder 2004). was originally described from the Middle Miocene (late Some (if not all) specimens previously described from Burdigalian/Langhian) of France and is very common in the Kirchberg Fm. and overlying OSM as Radix socialis coeval deposits of Central Europe (Harzhauser et al. 2014). dilatata by Schlickum (1964, 1966) and Reichenbacher Stagnicola praebouilleti Schlickum, 1970a, b, c, d (1989, 1993) probably belong to Radix socialis (von Zie- Figure 5c ten, 1830), identifiable by its diagnostic proportionally very minute spire (when compared to the large body whorl; e.g., 1964 Stagnicola (Stagnicola?) bouilleti – Schlickum: p. 14, Salvador and Rasser 2014). pl. 2, fig. 32 [non Michaud]. *1970d Stagnicola praebouilleti Schlickum: p. 92, Superfamily Planorboidea Rafinesque, 1815 figs. 12–17. Family Planorbidae Rafinesque, 1815 1973 Stagnicola (Stagnicola) praebouilleti – Steininger Genus Ferrissia Walker, 1903 et al.: p. 451, pl. 9, fig. 10. Ferrissia cf. wittmanni (Schlickum, 1964) 1993 Stagnicola praebouilleti – Reichenbacher: Table 13. Figure 5e 2005 Stagnicola praebouilleti – Kowalke and Reichen- Occurrence OBM (Grimmelfingen Fm.): Buttenwiesen. bacher: p. 631, figs. 9.8–9.9. OBM (Kirchberg Fm.): Buttenwiesen, Gu¨nzburg, Lauin- 2014 Stagnicola praebouilleti – Neubauer et al. 2014: gen, Offingen. supplementary material 1. Remarks These cap-shaped and elliptical shells, with a Occurrence OBM (Grimmelfingen Fm.): Buttenwiesen. longitudinally central apex, compare well to F. wittmanni,a OBM (Kirchberg Fm.): Buttenwiesen. species originally described as Ancylus wittmanni from the Remarks The forms known as S. praebouilleti, originally uppermost ‘‘Rzehakia Beds’’ (previously known as ‘‘On- described from the Bavarian ‘‘Oncophora Beds’’ (now cophora Beds’’; see also Kowalke and Reichenbacher 2005: ‘‘Rzehakia Beds’’, OBM), can be distinguished from S. p. 631, figs. 9.10–9.12). The species is well known from the armaniacensis by its slender shell and taller spire, deeper Kirchberg Fm. (Schlickum 1966; Reichenbacher 1989, suture and by usually having at least one additional half whorl. 1993) and its stratigraphic equivalents in upper Bavaria (Schlickum 1970a, b, c, d). Furthermore, F. wittmanni occurs Genus Lymnaea Lamarck, 1799 in the middle Miocene (Sarmatian) of Austria and Hungary Lymnaea dilatata (Noulet, 1854) (Harzhauser and Kowalke 2002). Nevertheless, the typical Figure 5d.

123 482 R. B. Salvador et al. axial sculpture (Kowalke and Reichenbacher 2005) could 2004 Gyraulus sp. A – Reichenbacher et al.: p. 76, pl. 1, not be observed in the present specimens, preventing a more figs. 4–5. definitive identification. 2004 Gyraulus sp. C – Reichenbacher et al.: p. 76, pl. 1, fig. 7. Genus Gyraulus Charpentier, 1837 2005 Gyraulus applanatus – Kowalke and Reichenbacher: Gyraulus albertanus (Clessin, 1877) p. 631, figs. 9.1–9.3. Figure 5f, g 2006 Gyraulus applanatus –Ko´kay: p. 56, pl. 19, *1877 Planorbis albertanus Clessin: p. 40. Figs. 13–14. 1923 Gyraulus (Gyraulus) albertanus – Wenz: p. 1541. 2006 Gyraulus trochiformis dealbatus –Ko´kay: p. 57, pl. 1972 Gyraulus (Gyraulus) albertanus – Gall: 16. 19, fig. 15. 2004 Gyraulus sp. B – Reichenbacher et al.: p. 76, pl. 1, 2014 Gyraulus applanatus – Neubauer et al.: supplemen- fig. 6. tary material 1. 2009 Menetus (Dilatata) albertanus – Moser et al.: p. 46. 2014 Gyraulus dealbatus – Neubauer et al.: supplementary 2014 Gyraulus albertanus – Neubauer et al.: supplemen- material 1. tary material 1. 2014 Gyraulus dealbatus – Salvador and Rasser: p. 192, 2014 Gyraulus albertanus – Salvador and Rasser: p. 191, figs. 16–23. figs. 12–15. Occurrence USM: Gundremmingen, Offingen. OBM Occurrence OBM (Grimmelfingen Fm.): Offingen. OBM (Grimmelfingen Fm.): Buttenwiesen, Offingen. OBM (Kirchberg Fm.): Holzheim, Offingen. OSM: Offingen. (Kirchberg Fm.): Buttenwiesen, Dillingen, Druisheim, ¨ ¨ Remarks This species is distinguished from its coeval Glott, Gunzburg, Hamlar 1, Holzheim, Lauingen, Offingen. ¨ congeners by its slightly larger size, more rounded shell OSM: Gempfing, Glott. profile, quickly growing whorls and large round aperture. It Remarks The present specimens show the common was originally described from Undorf (MN 5, SE Ger- conchological variation reported for G. applanatus in many; Clessin 1877) and is additionally known from several German sites, including the Kirchberg Fm. and Adelschlag (MN 5, SE Germany; Reichenbacher et al. OSM (Gottschick and Wenz 1916; Kowalke and 2004,asGyraulus sp.) and several other OSM localities Reichenbacher 2005; Salvador and Rasser 2014): two (Salvador and Rasser 2014). To our knowledge, this is the extremes, the typical G. applanatus and the typical G. first record of G. albertanus for the Grimmelfingen and dealbatus, with numerous intermediate forms. The typical Kirchberg Formations and thus, the earliest for the species. G. applanatus has a flattened shell, regularly growing Gyraulus applanatus (Thoma¨, 1845) whorls, more closely packed together, with a well-marked Figure 5h, i angulation on the laterobasal portion of the body whorl, and a smaller aperture, shaped as an arrowhead. The other *1845 Planorbis applanatus Thoma¨: p. 150. extreme, commonly called G. dealbatus in the literature, *1851 Planorbis dealbatus Braun: p. 1134. has a more rounded shell profile, with rapidly growing 1923 Gyraulus (Gyraulus) trochiformis applanatus – whorls, a faint angulation on its median portion and a Wenz: p. 1579. larger aperture. This variation, linked by numerous 1923 Gyraulus (Gyraulus) trochiformis dealbatus – Wenz: intermediate forms, led Kowalke and Reichenbacher p. 1591. (2005) to synonymize the two species, a decision which is 1964 Gyraulus trochiformis dealbatus – Schlickum: p. 15, followed here. pl. 2, fig. 35. 1970a Gyraulus trochiformis applanatus – Schlickum: Genus Planorbarius Dume´ril, 1806 p. 148, pl. 10, fig. 6. Planorbarius mantelli (Dunker, 1848) 1970c Gyraulus trochiformis applanatus – Schlickum: Figure 5j. p. 180. *1848 Planorbis Mantelli Dunker: p. 159, pl. 21, 1973 Gyraulus trochiformis dealbatus – Steininger et al.: figs. 27–29. p. 451, pl. 9, fig. 11a–b. 1966 Planorbarius cornu – Schlickum: p. 326, pl. 13, 1989 Gyraulus trochiformis dealbatus – Reichenbacher: fig. 27. p. 172, pl. 1, fig. 11. 1970a Planorbarius cornu – Schlickum: p. 149, pl. 10, 1993 Gyraulus trochiformis dealbatus – Reichenbacher: fig. 7. Tables 6–7, 13. 1989 Planorbarius cornu – Reichenbacher: p. 172, pl. 1, 1995 Gyraulus dealbatus – Kadolsky: p. 40, fig. 47. Fig. 10. 2004 Gyraulus dealbatus – Binder: p. 193, pl. 2, figs. 1a–c. 1993 Planorbarius cornu – Reichenbacher: Tables 6–7, 13.

123 Early Miocene continental gastropods from new localities of the Molasse Basin in southern… 483

2004 Planorbarius cornu – Binder: p. 193, pl. 2, figs. 2–3. site of Sandelzhausen (SE Germany; Salvador, 2015) and 2004 Planorbarius sp. – Reichenbacher et al.: p. 76, pl. 1, also known from the MN 5 locality of Riedensheim (Sal- figs. 1–3. vador et al. unpublished data). The present specimen has 2006 Planorbarius cornu cornu –Ko´kay: p. 58, pl. 20, weaker apertural barriers (especially the palatal tooth) fig. 6. when compared to C. galli, but intraspecific variation in 2009 Planorbarius cornu cornu –Bo¨ttcher et al.: p. 239, such features is well known in the genus (Strauch 1977; figs. 2.4–2.6. Stworzewicz 1999). The most characteristic feature of this 2014 Planorbarius mantelli – Harzhauser et al.: p. 15, pl. 3, species is the simple internal lamella (Salvador 2015). figs. 5, 7–13, 15–16. Nevertheless, to observe this feature, the specimen would 2014 Planorbarius mantelli – Neubauer et al., 2014: sup- have to be broken, which we opted not to do. plementary material 1. The externally similar species C. nouleti Bourguignat, 2014 Planorbarius cornu – Salvador and Rasser: p. 193, 1857 (and its likely synonym C. gibbum Sandberger, 1875; figs. 26–28. Strauch 1977; Harzhauser et al. 2014) differs from C. galli mainly by having a sinuous internal lamella (Salvador Occurrence USM: Gundremmingen. OBM (Grimmelfin- 2015). Other useful diagnostic characters of C. galli are gen Fm.): Buttenwiesen. OBM (Kirchberg Fm.): Butten- usually a more slender and smaller shell, a more defined wiesen, Dillingen, Druisheim, Glo¨tt, Gu¨nzburg, Holzheim, and spaced axial sculpture and a more angular shape of the Lauingen, Offingen. OSM: Buttenwiesen, Dillingen, peristome (Salvador 2015). Holzheim, Offingen. If the present identification is later confirmed, this would Remarks Only the first whorls and fragments of later be the earliest known record of C. galli. Furthermore, whorls are preserved. The shell size, shape and unique previous reports of C. nouleti from the uppermost parts of sculpture (protoconch of ca. 1 whorl sculptured by spiral the Kirchberg Fm. and ‘‘Rzehakia Beds’’ (previously lines of regularly organized circular pits, transitioning known as ‘‘Oncophora Beds’’; Schlickum 1964, 1966) abruptly to a teleoconch sculptured by spiral striae on first could actually represent C. galli. ca. 1 whorl and smooth on the rest, except for growth Stylommatophora A. Schmidt, 1855 lines) enable the identification as P. mantelli. This species Superfamily Zonitoidea Mo¨rch, 1864 is known from the late Early Miocene to the Late Miocene Family Zonitidae Mo¨rch, 1864 of Central Europe (Harzhauser et al. 2014). Specimens of Genus Archaeozonites Sandberger, 1872 P. mantelli from these localities and ages are usually Archaeozonites sp. referred to as P. cornu (Brongniart, 1810), a form from the Figure 5l Late Oligocene to Early Miocene. These hard to diagnose species are part of a complex that is in urgent need of Occurrence OBM (Grimmelfingen Fm.): Buttenwiesen. revision. OBM (Kirchberg Fm.): Offingen. Eupulmonata Haszprunar and Huber, 1990 Remarks Only fragmentary shell apexes remain, but the Superfamily Ellobioidea Pfeiffer, 1854 combination of its helicoid-like size and shape, flattened Family Carychiidae Jeffreys, 1830 first whorls and the characteristic teleoconch sculpture Genus Carychium O.F. Mu¨ller, 1773 (well-marked prosocline ribs) allows for the identification Carychium cf. galli Salvador, 2015 as Archaeozonites, a common member of land snail Figure 5k. assemblages in the Central European Oligocene and Mio- cene (e.g., Kadolsky 2008b; Salvador and Rasser 2014). Occurrence OBM (Grimmelfingen Fm.): Buttenwiesen. Superfamily Helicoidea Rafinesque, 1815 Remarks Thesinglespecimenavailableshowsamin- Family Helicidae Rafinesque, 1815 ute pupiform to fusiform shell, with a slightly acumi- Genus Megalotachea Pfeffer, 1930 nated spire (but with blunt apex) and its greatest width Megalotachea cf. silvana (Klein, 1853) on the body whorl. The dome-shaped protoconch (ca. Figure 5m–o 1 whorl) transitions abruptly to a teleoconch sculptured by fine prosocline axial riblets. The trapezoid Occurrence OBM (Grimmelfingen Fm.): Buttenwiesen. aperture has a faint parietal callus, a strong oblique OBM (Kirchberg Fm.): Glo¨tt, Holzheim, Lauingen, columellar lamella, a narrow parietal lamella and a very Offingen. OSM: Dillingen, Gempfing, Glo¨tt, Holzheim, faint median palatal tooth. Offingen. The present specimen compares well to the overall Remarks The specimens from the OSM layers are very morphology of C. galli, a species described from the OSM reminiscent of M. silvana, a common species from the

123 484 R. B. Salvador et al.

Silvana beds (previously classified in the genus Cepaea wide umbilicus) seem to point to the genus Discus, well- Held, 1838; e.g., Schlickum 1976; Reichenbacher 1989). known from the German Miocene. Unfortunately, the poor Nevertheless, due to their larger size and deformation due preservation prevents a precise identification, since the to sub-optimal preservation, we prefer here a more cautious characteristic sculpture pattern was not preserved. The classification. Moreover, the specimens from the OBM present specimen could represent Discus pleuradrus layers comprise fragmentary early whorls and may belong (Bourguignat, 1881), originally described from Sansan to another species. (MN 6; France), but known from the entire Miocene of Central and Western Europe (Bo¨ttcher et al. 2009). How- Vestiges of a color pattern can be seen under UV light, ever, as pointed out by previous authors, early and late consisting of three spiral bands on the whorl surface Miocene records could represent different species and, above the faint angulation and two additional spiral bands thus, D. pleuradrus would be restricted to the middle below it. Miocene (Moser et al. 2009; Harzhauser et al. 2014). Genus Palaeotachea Jooss, 1912 Palaeotachea sp Figure 5p–r Discussion Occurrence USM: Dillingen, Gundremmingen, Offingen. Continental mollusks, especially land snails, often show on Remarks Due to the poor preservation of the present the generic level preference for one type of habitat (Barker specimens, they cannot be identified beyond genus level. 2001; Cook 2001; Pearce and O¨ rstan 2006). Therefore, These specimens, all from the USM, are much smaller than ecological data from recent genera are often used as a the M. cf. silvana found in the OSM layers. They also guide for paleoecological inferences of congeneric fossil apparently have a proportionately much smaller aperture species (e.g., Albesa et al. 1997; Moser et al. 2009). Below, and a greater constriction in the body whorl immediately we present an analysis regarding each major stratigraphical preceding the aperture. By the shells’ size and overall unit; Table 1 lists the occurrence of each species in these shape, as well as the age of the sediments, the present units. Since the sites are all close to each other (Fig. 1), specimens likely belong to P. subsulcosa (Thoma¨, 1845). they are treated together; nevertheless, some localities (or group of localities) receive further comments separately Family Hygromiidae Tryon, 1866 when appropriate. Genus Leucochroopsis Boettger, 1909 Leucochroopsis sp. USM The molluscan fauna recovered from the USM Figure 5s. layers is rather poor, which could be a reflection of the low amounts of fossil-bearing samples available. As expected, Occurrence OBM (Grimmelfingen Fm.): Buttenwiesen. it is composed mainly of freshwater snails (Table 1). The OBM (Kirchberg Fm.): Offingen. habitats preferred by their living congeners are richly Remarks The material can be identified as Leucochroopsis vegetated standing or slowly flowing waters, sometimes by its size, depressed spire, whorls regularly increasing in even temporary water bodies (Welter-Schultes 2012). The size and characteristic sculpture: protoconch (ca. 1 whorl) land snails recovered are ?Discus sp., whose rather sculptured by fine parallel striae, transitioning abruptly to a uncertain identity precludes the use in a paleoenviron- teleoconch sculptured by fine, regularly distributed, well- mental analysis, and Palaeotachea sp., which should thrive marked papillae, which, together with the growth lines, give in a rather broad range of habitats (hot humid temperate to the impression of prosocline axial striae. Identification sub-tropical, in reasonably well-vegetated areas). beyond genus level is not possible, but there are records of L. Only a few previous studies have dealt with the pale- francofurtana (Wenz, 1919) for the OBM (Schlickum 1964; oecological interpretation of the USM (e.g., Reichenbacher Reichenbacher 1989). and Weidmann 1992; Reichenbacher 1996; Scha¨fer 2005, Superfamily Punctoidea Morse, 1864 2011; Weidmann et al. 2014). All these studies were based Family Discidae Thiele, 1931 on fossil assemblages found in the westernmost Molasse Genus Discus Fitzinger, 1833 Basin, i.e., in western Switzerland and Haute Savoie, and ?Discus sp gastropods have not been considered. However, the sug- Figure 5t, u gestion of heterogenous lacustrine and terrestrial habitats Occurrence USM: Gundremmingen. with some periods of drought from time to time indicated by the above-mentioned studies (based on charophytes, Remarks The small size and overall shape of the shell ostracods, fish otoliths and mammal remains) fits with the (discoid, with low spire, whorls regularly growing, and interpretation presented here based on the gastropods. 123 Early Miocene continental gastropods from new localities of the Molasse Basin in southern… 485

Table 1 List of all species found in the present material and their to the conchological identical genus Heleobia Stimpson, general stratigraphical occurrence (considering all localities together) 1865 (family Cochliopidae Tryon, 1866), which is more Species OSM OBM USM typically found in freshwater (although also occurring in brackish environments). uKF lKF GF Almost all previous reports on the paleoenvironment of Neritimorpha the Grimmelfingen Fm. refer to marine or brackish faunal Theodoxus cyrtocelis XXXXX elements found in the base of this formation (Reichen- Theodoxus obtusangula X bacher et al. 1998, 2013; Sach and Heizmann 2001). Only Caenogastropoda Viviparus and the bivalve Rzehakia Korobkov, 1954 have Melanopsis impressa XX already been reported from the upper parts of the Grim- Tinnyea lauraea XX melfingen Fm. (Reichenbacher 1989, 1993). The freshwa- Bithynia glabra XXXXXter and terrestrial gastropod species described here are, Ctyrokya conoidea ?XX thus, the first record for the Grimmelfingen Fm. The gas- Hydrobia semiconvexa XX tropods indicate that the environment may have been much Nematurella bavarica ?XXX less brackish and estuarine than previously thought. The ?Truncatella sp. X environment of these layers could be freshwater or of very Viviparus cf. suevicus X low salinity levels. Heterobranchia On land, the presence of the hygrophilous land snail ?Valvata sp. X X Carychium is a fair indicative of humid and usually well- Pulmonata vegetated areas near a water body (Welter-Schultes 2012). Stagnicola armaniacensis XXXXXA single specimen of Carychium cf. galli was found in Stagnicola praebouilleti XX Buttenwiesen, but the fragile and diminutive shells of the Lymnaea dilatata XX XXgenus makes preservation and/or recovery difficult (and Ferrissia cf. wittmanni XX this locality is the one with the better-preserved speci- Gyraulus albertanus XX X mens). Archaeozonites and Leucochroopsis are also con- Gyraulus applanatus XXXXXsidered indicatives of humid forests (Lueger 1981). Planorbarius mantelli XXXXX OBM (lower Kirchberg Fm.) The molluscan fauna of the Carychium cf. galli X lower Kirchberg Fm. is very different to that of the Grim- Archaeozonites sp. X melfingen Fm., likely indicating changes in the environmental Megalotachea cf. silvana XX X conditions. These beds ‘‘lose’’ much of their pulmonate Palaeotachea sp. X aquatic fauna and show the richest operculate snail fauna (i.e., Leucochroopsis sp. X non-pulmonates) of all layers studied here (Table 1). ?Discus sp. X X Theodoxus (which occurs in considerable numbers in the The presence of Nematurella bavarica and Ctyrokya conoidea in the OSM is interpreted as reworked material from older OBM layers (see Kirchberg Fm.) and Viviparus are typical of freshwater ‘‘Discussion’’) environments, although a few recent species can also be GF Grimmelfingen Formation, lKF lower Kirchberg Formation; OBM found in low salinity waters (Bandel, 2001; Gloër and Upper Brackish Molasse, OSM Upper Freshwater Molasse; uKF Meier-Brook 2003; Zettler et al. 2004; Bunje 2005; Welter- upper Kirchberg Formation, USM Lower Freshwater Molasse Schultes 2012). Tinnyea lauraea have been found in both freshwater and oligohaline sediments (Kadolsky 1995; OBM (Grimmelfingen Fm.) These beds have fauna more Kowalke 2004;Ko´kay 2006). The more typical brackish typical of freshwater environments, as indicated by the fauna of the lower Kirchberg Fm. includes the hydrobiids richness and abundance of freshwater species: Bithynia, (although with reservations, as explained above), possibly Theodoxus and the basommatophoran pulmonates ?Truncatella sp. and several brackish water bivalves (not (Table 1; despite being preferentially freshwater species, studied here, but known from previous reports, e.g., Sch- some recent congeners of the operculate snails can with- lickum 1963; Reichenbacher 1989). stand slightly brackish waters). The hydrobiids, despite The environment of the lower Kirchberg Fm. is thus being considered more typical brackish water snails interpreted as low brackish to brackish (with a salinity level (especially the genus Nematurella), are also commonly still allowing the scarce pulmonates, Bithynia, Theodoxus, found in fully freshwater environments (e.g., Welter- Viviparus and ?Valvata to thrive). This assumption is Schultes 2012). Kadolsky (2008a) even raised the possi- consistent with the suggestion of a brackish environment bility that some Miocene hydrobiids could actually belong based on the fish fauna (Reichenbacher 1993).

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Melanopsis impressa, in particular, is an index for Table 2 List of the freshwater species found in the northeastern strong fluvial influx (Harzhauser and Kowalke 2001). The localities (NE; Druisheim, Gempfing and Hamlar 1; see also Fig. 1) fragmented nature of the shells of M. impressa from Agapi and the remaining southwestern localities (SW) (early Miocene, Greece) was deemed by Harzhauser and Species SW NE Kowalke (2001) as a sign that the sedimentation area was uKF lKF uKF lKF far from the more energetic environments where the species usually occurs. This is similar to what is seen in the Theodoxus cyrtocelis XXXX present specimens, although breakage could have happened Theodoxus obtusangula X during sample processing. Melanopsis impressa XXXX A single specimen of ?Truncatella sp. was found in the Tinnyea lauraea XXX locality ‘‘Hamlar 1’’. The recent Truncatella subcylindrica Bithynia glabra XXXX (Linnaeus, 1767) lives in the marine supralittoral, usually Ctyrokya conoidea XXXX out of the water on algae or burrowed in the sand and it can Hydrobia semiconvexa XX survive through long periods of submersion (Welter- Nematurella bavarica XXXX Schultes 2012). Since only a single specimen of dubious ?Truncatella sp. X identification was found, its use in a paleoecological Viviparus cf. suevicus X analysis must remain tentative. Its poor preservation could ?Valvata sp. X X indicate that it was carried from the littoral by some cur- Stagnicola armaniacensis XX rents. Nevertheless, a unique occurrence of a foraminifera Stagnicola praebouilleti X layer intercalated into the ‘‘regular’’ Kirchberg Fm. (Re- Lymnaea dilatata XX ichenbacher et al. 2013: Fig. 5) was found in the locality Ferrissia cf. wittmanni X ‘‘Hamlar 1’’, which could indicate a rapid small-scale Gyraulus albertanus X transgression. Another (less likely) possibility is that this Gyraulus applanatus XXXX ?Truncatella sp. could respresent a fully terrestrial species, Planorbarius mantelli XX X a trait known to have evolved repeatedly in Truncatella and lKF lower Kirchberg Formation, uKF upper Kirchberg Formation other truncatellids (Rosenberg 1996). OBM (upper Kirchberg Fm.) Overall, the layers of the upper Kirchberg Fm. have a fauna of operculate snails local fluvial influx, while the NE localities would have similar to the lower levels, but, at the same time, have a retained slightly more brackish environments (similar to more diverse basommatophoran fauna. As in the Grim- the lower Kirchberg Fm.). melfingen Fm., this should indicate a fully or nearly Finally, one unusual occurrence noted in the present freshwater environment. A similar assumption has previ- material is the record of Ctyrokya conoidea for the ously been presented based on the microfossil assemblages upper Kirchberg Fm. of some localities (see Online and especially the fish otoliths, which were found to be Supplementary Appendix 1), since previous reports indicative for a mesohaline to oligohaline environment in restrict this species to the lower Kirchberg Fm. (e.g., the upper Kirchberg Fm. (Reichenbacher 1989, 1993). The Schlickum 1966; Reichenbacher 1989). This means that presence of the land snails Archaeozonites and Leu- C. conoidea survived the transition in some localities. cochroopsis would again indicate humid forests in the In Offingen, the species is only recorded from the most surrounding area. basal layers of the upper Kirchberg Fm. and, thus, managed to survive only for a brief time. However, in The freshwater snails of the northeasternmost boreholes other localities such as Buttenwiesen, C. conoidea (Druisheim, Gempfing and Hamlar 1; Fig. 1) seem to thrived for longer, disappearing only towards the end of present a different composition than the other localities. In the Kirchberg Fm. the lower Kirchberg Fm., the faunas are very similar (Table 2), except for the occurrence of two additional OSM In the OSM layers, most of the operculate snails species in the NE localities that are more commonly related disappear from the record, causing the diversity of aquatic to freshwater: ?Valvata sp. and Viviparus cf. suevicus.In species to greatly decrease. This is consistent with other the upper Kirchberg Fm., however, the NE localities show faunas from German localities (e.g., Salvador and Rasser a much less diverse typical freshwater fauna, with only two 2014), which can be very abundant in sheer number of pulmonates recorded (Table 2). The other localities have freshwater specimens, but rather poor in richness of fresh- records of seven pulmonate species and also ?Valvata sp. water species (composed mainly of pulmonates). The same As such, it is plausible that the SW localities possessed pattern is seen in the otolith-based fish fauna from the fully or nearly freshwater environments, probably due to lowermost OSM: few freshwater and/or euryhaline species 123 Early Miocene continental gastropods from new localities of the Molasse Basin in southern… 487 are present, often represented by numerous specimens (e.g., Bo¨hme, M. 2010. Ectothermic vertebrates (Actinopterygii, Allocau- Reichenbacher et al. 2004; Reichenbacher and Prieto 2006). data, Urodela, Anura, Crocodylia, Squamata) from the Miocene of Sandelzhausen (Germany, Bavaria) and their implications for As in the USM, discussed above, most of the species environment reconstruction and palaeoclimate. Palaöntologische present in the OSM prefer richly vegetated standing or Zeitschrift 84: 3–41. Bo¨ttcher, R., E.P.J. Heizmann, M.W. Rasser, and R. Ziegler. 2009. slow-moving waters. The possibly brackish water species Biostratigraphy and palaeoecology of a Middle Miocene Nematurella bavarica and Ctyrokya conoidea recorded (Karpathian, MN 5) fauna from the northern margin of the from basal layers of some OSM localities (see Online North Alpine Foreland Basin (Oggenhausen 2, SW’ Germany). Supplementary Appendix 1) very likely represent reworked Neues Jahrbuch fu¨r Geologie und Palaöntologie Abhandlungen 254(1–2): 237–260. material from older OBM layers. Bouchet, P., J.P. Rocroi, J. Fry´da, B. Hausdorf, W. Ponder, A´ . Valde´s, and A. Waren. 2005. Classification and nomenclator of gas- Acknowledgments We are deeply grateful to Karin Wolf-Sch- tropod families. Malacologia 47: 1–397. wenninger (SMNS) for the SEM images; to Thomas A. Neubauer Bourguignat, M.J.R. 1857. Ameńite´s Malacologiques, LXIV. Du (NHMW) for the photos and information on the syntypes of H. genre Carychium. Revue et Magasin de Zoologie (2) 9(5): frauenfeldi; to Dietrich Kadolsky for pointing out the poper spelling 209–232. of the specific epithet of Theodoxus obtusangula; to Olaf Ho¨ltke Bourguignat, M.J.R. 1881. Histoire Malacologique de la Colline de (SMNS) for the help with the helicid snails; to Simon Schneider, T.A. Sansan. Annales des Hautes E´ tudes, Sciences Naturelles 22(3): Neubauer and Mike Reich for their suggestions, which greatly 1–175. improved the present manuscript. This work was partly supported by Braun, A. 1851. Die fossile Fauna des Mainzer Beckens. Wirbellose a doctorate grant from CNPq (Conselho Nacional de Desenvolvi- Thiere. In Handbuch der Geognosie, 2nd ed., ed., F.A. Walchner, mento Cientı´fico e Tecnolo´gico, Brazil) to R.B.S (Process #245575/ 1112–1141. Karlsruhe. 2012-0). Brongniart, M. 1810. 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