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The Fossil Pulmonate Snails of Sandelzhausen (Early/Middle Miocene, Germany): Ellobiidae, Pupilloidea, and Clausilioidea

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The Fossil Pulmonate Snails of Sandelzhausen (Early/Middle Miocene, Germany): Ellobiidae, Pupilloidea, and Clausilioidea The fossil pulmonate snails of Sandelzhausen (Early/Middle Miocene, Germany): Ellobiidae, Pupilloidea, and Clausilioidea Rodrigo Brincalepe Salvador Paläontologische Zeitschrift Scientific Contributions to Palaeontology ISSN 0031-0220 Volume 89 Number 1 Paläontol Z (2015) 89:37-50 DOI 10.1007/s12542-013-0210-4 1 23 Your article is protected by copyright and all rights are held exclusively by Springer- Verlag Berlin Heidelberg. This e-offprint is for personal use only and shall not be self- archived in electronic repositories. If you wish to self-archive your article, please use the accepted manuscript version for posting on your own website. You may further deposit the accepted manuscript version in any repository, provided it is only made publicly available 12 months after official publication or later and provided acknowledgement is given to the original source of publication and a link is inserted to the published article on Springer's website. The link must be accompanied by the following text: "The final publication is available at link.springer.com”. 1 23 Author's personal copy Pala¨ontol Z (2015) 89:37–50 DOI 10.1007/s12542-013-0210-4 RESEARCH PAPER The fossil pulmonate snails of Sandelzhausen (Early/Middle Miocene, Germany): Ellobiidae, Pupilloidea, and Clausilioidea Rodrigo Brincalepe Salvador Received: 19 March 2013 / Accepted: 23 September 2013 / Published online: 16 October 2013 Ó Springer-Verlag Berlin Heidelberg 2013 Abstract Sandelzhausen is an Early/Middle Miocene Mollusken, nahezu ausschließlich pulmonate Gastropoden, (Mammal Neogene zone MN5) fossil site near Mainburg, S die bei Ausgrabungen geborgen wurden, aber nur wenig Germany, that despite its small size harbors a rich fossil wissenschaftliche Beachtung fanden. Vorliegende Studie record. Thousands of fossil continental mollusks, almost stellt den ersten Teil einer formalen taxonomischen Bear- exclusively pulmonate snails, were recovered during the beitung der fossilen Pulmonaten von Sandelzhausen dar. excavations, but did not receive much attention from Sie behandelt die U¨ berfamilien Ellobioidea, Pupilloidea researchers. Here, the first part of a formal taxonomic und Clausilioidea, einschließlich der Beschreibung einer treatment of Sandelzhausen’s fossil pulmonates is pre- neuen Art. Folgende Arten wurden gefunden: Carychium sented, dealing with the superfamilies Ellobioidea, Pupil- eumicron and Carychium galli sp. nov. (Ellobiidae); Gra- loidea, and Clausilioidea, and including the description of a naria cf. grossecostata und Granaria sp. (Chondrinidae); new species. The following species were found in the Gastrocopta acuminata und Gastrocopta nouletiana material: Carychium eumicron and Carychium galli sp. (Gastrocoptidae); ?Pyramidula sp. (Pyramidulidae); nov. (Ellobiidae); Granaria cf. grossecostata and Granaria Strobilops sp. (Strobilopsidae); Vallonia lepida (Vallonii- sp. (Chondrinidae); Gastrocopta acuminata and Gastro- dae); Vertigo callosa (Vertiginidae); Pseudidyla moer- copta nouletiana (Gastrocoptidae); ?Pyramidula sp. (Pyra- singensis (Clausiliidae); Triptychia sp. (Filholiidae). midulidae); Strobilops sp. (Strobilopsidae); Vallonia lepida (Valloniidae); Vertigo callosa (Vertiginidae); Pseudidyla Schlu¨sselwo¨rter Carychium galli nov. spec. Á moersingensis (Clausiliidae); Triptychia sp. (Filholiidae). Gastropoda Á Mammal Neogene Zone MN5 Á Pulmonata Á Stylommatophora Keywords Carychium galli new species Á Gastropoda Á MN5 European Mammal Neogene zone Á Pulmonata Á Stylommatophora Introduction Kurzfassung Sandelzhausen ist eine mittelmioza¨ne The Sandelzhausen fossil site is a Fossil-Lagersta¨tte and (Sa¨ugerzone MN5) Fossilfundstelle nahe Mainburg in one of the most important continental sites in Europe Su¨ddeutschland. Trotz Ihrer geringen geographischen (Moser et al. 2009a). Besides some plant remains, a fauna Ausdehnung bietet sie eine reiche Fauna an kontinentalen of more than 200 metazoan species (and 50,000 specimens) was collected, including mollusks, ostracods, and repre- sentatives of all vertebrate classes, although the mammals R. B. Salvador (&) Staatliches Museum fu¨r Naturkunde Stuttgart, have received the most attention (Moser et al. 2009a, and Stuttgart, Germany references therein). Gastropods were among the first fossils e-mail: [email protected] found in Sandelzhausen, and thousands of specimens were collected before the site was closed in the last decade. Only R. B. Salvador Mathematisch-Naturwissenschaftliche Fakulta¨t, Eberhard Karls three works dealt specifically with these mollusks. Gall Universita¨tTu¨bingen, Tu¨bingen, Germany (1972) identified 49 gastropod and two bivalve species, but 123 Author's personal copy 38 R. B. Salvador based his work heavily on faunas such as Sansan, in Pala¨ontologie und Geologie (BSPG; Munich, Germany) France, that are now known to be younger (Moser et al. under the record number BSPG 1959 II. All available 2009b). Gall (1973) dealt with aspects of the taphonomy specimens of the three superfamilies were analyzed for this and preservation of the larger shells. Moser et al. (2009b) work; a list of the examined material follows each species dealt with paleoecological questions, and thus presented description. All specimens in a good state of preservation only a preliminary classification of the mollusks. The were measured either with digital calipers or with the aid of present study is the first part of a taxonomic treatment of computer software. Abbreviations used with shell mea- the fossil mollusks from Sandelzhausen, dealing with the surements: H = shell length; D = shell greatest width; superfamilies Ellobioidea, Pupilloidea, and Clausilioidea S = spire length (excluding aperture); S0 = spire length (12 species in total). Further works describing the (excluding body whorl); h = aperture height; d = aperture remaining fauna will soon follow (e.g., Salvador 2013). width. The spire angle used here is the ‘‘mean spire angle’’ sensu Moore (1960). Selected specimens were examined by scanning electronic microscopy (SEM) in the Staatli- Geological setting ches Museum fu¨r Naturkunde Stuttgart (SMNS; Stuttgart, Germany). Since only two works have attempted to iden- The Sandelzhausen fossil site was located in the vicinity of tify the mollusks from Sandelzhausen (Gall 1972; Moser the city of Mainburg, 60 km north of Munich, in the Mo- et al. 2009b), the previous identification given by these lasse Basin (Molassebecken) of southern Germany, which authors was included here in order to facilitate future harbors the ‘‘formation’’ known as the Upper Freshwater correspondence among the works. Molasse (Ober Su¨ßwassermolasse, OSM; Moser et al. The excavation site in Sandelzhausen was divided into a 2009a). The fossils from Sandelzhausen derive from a square grid, so a good part of the material has precise data member of the OSM called No¨rdlicher Vollschotter, about where it was found. A portion of the material shows composed primarily of marl and gravel (Moser et al. data on the original height within the section and can thus 2009a). The age of these fossiliferous deposits was estab- be attributed to the layers published by Fahlbusch and Gall lished by stratigraphic, biostratigraphic, and magneto- (1970) and Moser et al. (2009b). However, a good portion stratigraphic correlations: the Early/Middle Miocene of the material, despite having the original height in the Burdigalian/Langhian boundary, early middle MN5 Euro- sediment recorded, cannot be unmistakably attributed to pean Mammal Neogene zone (*16.47–16.27 Ma; Moser the layers, since the given measurements are not precise et al. 2009a). enough (Markus Moser, personal communication). More- The facies types of the deposits from Sandelzhausen over, many specimens completely lack locality data, while were studied by Fahlbusch and Gall (1970) and Moser et al. some have labels that cannot be traced back to a precise (2009a). From bottom to top: layer A: marly gravels, point within the excavation site, such as the samples sometimes cemented by carbonates; fossil content rare and labeled ‘‘Grube Bergmaier’’ (taken during the earliest test limited to robust skeletal parts; layer B: gravel-rich marl in excavations by Richard Dehm; Moser et al. 2009b) and which the size and number of pebbles diminish upwards, ‘‘Testprofil 1966’’ (a test section preceding the start of the with intercalated sand horizons; origin of most macrover- main excavations). As such, only the samples that can be tebrate fossils; layer C: fossil-rich marl divided into three attributed to distinct layers were taken into account here to smaller layers (C1, C2, and C3) by a black, organic rich determine the stratigraphical range of each species. layer (C2); layer D: marl (mainly silt) with few pebbles and diffuse carbonates and carbonatic nodules; rich in Systematics fossils, many in an excellent state of preservation due to a less intense compaction; layer E: silty clays with micro- Superfamily Ellobioidea Pfeiffer, 1854 vertebrate fossils; Layer F: laminite with alternating light Family Ellobiidae Pfeiffer, 1854 and dark bands, carbonate concretions and desiccation Genus Carychium O.F. Mu¨ller, 1773 cracks; no fossils. Fossil mollusks can be found in layers A to D; a more detailed description of the lithology was given Carychium (Carychiella) eumicron Bourguignat, 1857 by Moser et al. (2009a). (Fig. 1) Carychium eumicron Bourguignat, 1857: 253. Materials and methods Carychium eumicron peneckei: Wenz, 1923b: 1190; Gall, 1972: 14. All of the material from Sandelzhausen is housed
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