Fossil terrestrial and freshwater Gastropoda from the Eary ... · Rodrigo B. Salvador & Michael W. Rasser Staatliches Museum für Naturkunde Stuttgart, Rosenstein 1, 70191, Stuttgart,

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Archiv für Molluskenkunde 146 (2) 233–241 Frankfurt am Main, 20 Dec. 2017

© E. Schweizerbart’sche Verlagsbuchhandlung (Nägele u. Obermiller) and Senckenberg Gesellschaft für Naturforschung, 2017, ISSN 1869-0963DOI 10.1127/arch.moll/146/233-241

Fossil terrestrial and freshwater Gastropoda from the Early/Middle Miocene of Heuchlingen, Germany

Rodrigo B. Salvador & Michael W. Rasser

Staatliches Museum für Naturkunde Stuttgart, Rosenstein 1, 70191, Stuttgart, Germany. • Corresponding author: R.B. Salvador ([email protected]).

Abstract. The present study is a report of fossil terrestrial and freshwater gastropods from a new out-crop exposed in the 1990s in Heuchlingen (Baden-Württemberg, SW Germany). The fossils occur in the Silvana Beds of the Upper Freshwater Molasse (OSM), dating from the Early/Middle Miocene (Euro-pean Land Mammal Zone MN 5). Thirty gastropod species (6 aquatic and the remainder terrestrial) were found, belonging to the families Pomatiidae, Lymnaeidae, Planorbidae, Ellobiidae, Succineidae, Cochli-copidae, Chondrinidae, Vertiginidae, Gastrocoptidae, Clausiliidae, Ferussaciidae, Subulinidae, Discidae, Zonitidae, Agriolimacidae, Helicidae, Helicodontidae, Hygromiidae, and Trissexodontidae. These fossils are used as paleoecological proxies in an actualistic genus-level analysis to present a first attempt at an environmental reconstruction of the Heuchlingen locality. The gastropod fauna points to a paleoenviron-ment consisting of a shallow lake (or a shallow margin of a larger lake) surrounded by humid forests and scrublands. Literature records from Heuchlingen and the nearby locality Dettingen am Albuch, as well as the surviving voucher material, are also reviewed.

Key words. Caenogastropoda, Dettingen am Albuch, European Land Mammal Zone MN 5, Pulmonata, Silvana Beds, Upper Freshwater Molasse.

DOI. https://doi.org/10.1127/arch.moll/146/233-241

IntroductionThe Miocene Upper Freshwater Molasse (“Obere Süß-wassermolasse” in German; abbreviated OSM) has nu-merous fossiliferous outcrops in southern Germany (e.g., Sandberger 1870–1875, Wenz 1923–1930), some of which might be considered fossil Lagerstätten (Abdul-Aziz et al. 2008, 2010). The Silvana Beds are a unit of the OSM famous for its abundance of non-marine gastropod fossils, which are usually well preserved. These beds are named after the most commonly found fossil land snail, Palaeotachea silvana (Klein, 1853), and are assigned to the European Land Mammal Zone MN 5, which ranges from c. 15.8 to 13.8 Ma (Esu 1999, Berger et al. 2005).

Heuchlingen is a district of the Gerstetten municipal-ity, Baden-Württemberg, southwestern Germany (Fig. 1). Fossil outcrops from this region have scarcely ever been reported in the literature. There is a single work reporting OSM outcrops and fossil mollusks from Heuchlingen, by Kranz et al. (1924). That work presents a list of mol-lusks attributed to malacologist Carlo G.H. Jooss, from 3 outcrops of the Silvana Beds in the region of Heuch-lingen and nearby Dettingen am Albuch (another district

of Gerstetten). Jooss reported 23 gastropod species and fragments of bivalves from there, but the localities have never since been mentioned in the literature.

A new outcrop was exposed in Heuchlingen in the 1990s, during construction work. Sediment samples were collected and stored in the collection of the Staatliches Museum für Naturkunde Stuttgart (SMNS; Stuttgart, Germany). Here we present a concise report of the fossil terrestrial and freshwater gastropods from this outcrop. Moreover, as gastropods can be used as paleoecological

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Figure 1. Map showing the location of Heuchlingen, Ger-stetten municipality, Heidenheim district, Baden-Württemberg state (BW), Germany.


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Archiv für Molluskenkunde · 146 (2) 2017

proxies for environmental reconstructions (e.g., Ložek 1964, Clarke 1979, Fordinál 1996, Albesa et al. 1997, Esu & Ciangherotti 2004), we also present here a first attempt at this for the Heuchlingen fauna.

Geological setting. The outcrop in Heuchlingen was ex-posed in the mid-1990s during excavations for construction work; the precise location and nature of the excavation are now unknown. A large amount of sediment was collected in a single take by a Mr Heizmann in 1996 and donated to the SMNS. However, neither lithological nor sedimento-logical data were taken and no surviving field notes could be traced in the SMNS collections or archives. Colleagues working at the SMNS at that time also could not provide any additional information. The sediment from this out-crop was later screenwashed for mammalian fossils and deposited in the collection of the SMNS.

This outcrop is no longer available and, unfortunately, no other outcrops (including those explored by Jooss) are presently known. Therefore, only literature data are available regarding the geology of this locality. According to Kranz et al. (1924), the OSM Silvana Beds sediments in this area (including Dettingen am Albuch) consist of yellowish or greenish limestones that may reach up to 25 m in thickness, which could indicate a rather long-lived lacustrine system. The material studied by Jooss comes from these limestones and the surviving vouchers are housed in the SMNS.

Kranz et al. (1924) assigned the sediments from Heuch lingen to the Silvana Beds. The composition of the gastropod fauna, as discussed below, is typical of the Sil-vana Beds, especially in comparison with other outcrops from Baden-Württemberg, thus supporting a MN 5 age for these sediments. Furthermore, fossils of Amphicyo-nidae carnivores (SMNS 46057, 46058) from the same samples as the mollusks presented here are consistent

with other outcrops of MN 5–6 age in southern Germany (Nagel et al. 2009, Morlo et al. 2010, R. Ziegler & E. Heizmann personal communication 2016).

Materials and MethodsThe material is deposited in the collection of the SMNS. The lots containing gastropods are indicated in Table 1 for each species (additionally, a few ostracods were recovered, lots SMNS 107213, 107214). As the sediment had been screenwashed for mammalian fossils, larger gastropod shells were fragmented, so that only shell apexes of the larger snails were preserved. The smaller snails, however, are very well preserved.

All gastropod species found are illustrated (Figs 2–4); the images were obtained with a scanning electron microscope (SEM) at the SMNS. Preserved specimens from Jooss’s collection (vouchers from Kranz et al. 1924) were conventionally photographed (Fig. 5). Measurements were made with the software ImageJ (Rasband 2012) and are referred to by the following abbreviations: H = shell height (parallel to coiling axis); D = greatest shell width, perpendicular to H; h = greatest operculum length; d = greatest operculum width.

Results and Discussion

SystematicsThe gastropod species found in Heuchlingen are listed in Table 1, grouped into selected higher clades and arranged within them by families (classification follows Bouchet et al. 2005 and Nordsieck 2014). We will not

Table 1. List of molluscan species recorded from Heuchlingen, with the collection number of the lots deposited in the SMNS collection and paleoecological information, the relative abundance of each species in the sample, the preferred habitat(s) and the reference(s) for such data (when necessary, indicating the supposed most closely-related living genus). Abbreviations and symbols: HW = humid woods; MA = marsh, stagnant waters; RB = reed belt of water body; RM = rocky (calcareous) meadows; SL = scrublands; SW = slow moving waters; † = exclusively fossil genus; * = living congeners have habitat preferences that are too broad for meaningful paleoecological interpretation.

Species (and higher taxa) Figure(s) Material (SMNS)Paleoecology

Relative abundance Habitat Reference(s)

CAENOGASTROPODA

Family Pomatiidae

Pomatias conicus (Klein, 1853) 2A–C 107258, 107259 1.0% HW, SLKerney & Cameron (1979), Welter-

Schultes (2012)

PULMONATA: HYGROPHILA

Family Lymnaeidae

Radix socialis (Zieten, 1832) 2D 107300 3.0% SW Welter-Schultes (2012)

Family Planorbidae

Ferrissia deperdita (Desmarest, 1814) 2E 107233 <1.0% RB Glöer (2002)

Gyraulus applanatus (Thomä, 1845) 2F–J 107244 16.0% SW Welter-Schultes (2012)


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Salvador & Rasser · Miocene land and freshwater snails from Heuchlingen

Species (and higher taxa) Figure(s) Material (SMNS)Paleoecology

Relative abundance Habitat Reference(s)

Hippeutis subfontanus (Clessin, 1877) 2K 107245 <1.0% SW Welter-Schultes (2012)

Planorbarius mantelli (Dunker, 1848) 2L–M 107254, 107363 27.0% SW Welter-Schultes (2012)

Segmentina lartetii (Noulet, 1854) 2N–P 107238 2.0% MA Clark (2011), Welter-Schultes (2012)

PULMONATA: ELLOBIOIDEA

Family Ellobiidae

Carychium eumicrum Bourguignat, 1857 3M 107249 <1.0% * Welter-Schultes (2012)

Carychium galli Salvador, 2015 3N 107250 3.0% * Welter-Schultes (2012)

PULMONATA: STYLOMMATOPHORA

Family Succineidae

Oxyloma minima (Klein, 1853) 4F 107261 <1.0% RB Welter-Schultes (2012)

Family Cochlicopidae

Hypnophila loxostoma (Klein, 1853) 3F 107251 2.5% SL Welter-Schultes (2012)

Family Chondrinidae

Granaria sp. 3B 107253 2.5% RM Höltke & Rasser (2013)

Family Vertiginidae

Negulopsis lineolata (Sandberger, 1872) 4H 107243 <1.0% HWSalvador et al. (2016b)

[Negulus Boettger, 1889]

Vertigo angulifera Boettger, 1884 4I 107298 <1.0% * Welter-Schultes (2012)

Vertigo callosa (Reuss, 1849) 4J 107252 <1.0% * Welter-Schultes (2012)

Family Gastrocoptidae

Gastrocopta acuminata (Klein, 1846) 3P 107256 6.5% * Pilsbry (1948), Zilch (1959–1960)

Gastrocopta nouletiana (Dupuy, 1850) 3Q 107255 6.5% * Pilsbry (1948), Zilch (1959–1960)

Family Clausiliidae

Pseudidyla moersingensis (O. Boettger, 1877) 3C–E 107232 2.5% HW† Nordsieck (2007)

Family Ferussaciidae

Cecilioides aciculella (Sandberger, 1874) 3O 107248 <1.0% * Welter-Schultes (2012)

Family Subulinidae

Opeas minutum (Klein, 1853) 4E 107247, 107366 5.0% HW Zilch (1959–1960), Willig et al. (2013)

Family Discidae

Discus euglyphoides (Sandberger, 1872) 3G–I 107246 3.0% HW Welter-Schultes (2012)

Discus pleuradrus (Bourguignat, 1881) 3J–L 107242, 107364 3.5% HW Welter-Schultes (2012)

Family Zonitidae

Archaeozonites sp. 4K 107241 <1.0% HW† Lueger (1981)

Family Agriolimacidae

Deroceras sp. 3A 107240 <1.0% * Rowson et al. (2014)

Family Helicidae

Palaeotachea sp. 3R 107237 5.0% SL, HW† Welter-Schultes (2012) [Cepaea Held, 1838]

Pseudochloritis sp. 3S 107236 2.5% HW† Höltke & Rasser (2015)

Family Helicodontidae

Protodrepanostoma involutum (Thomä, 1845) 3T–U 107239 <1.0% HW† Cameron (1972), Maltz (2007) [Helicodonta Férussac, 1821]

Family Hygromiidae

Leucochroopsis kleinii (Klein, 1846) 4A–C 107215 3.0% HW† Lueger (1981)

Urticicola perchtae Salvador, 2013 4D 107299 <1.0% * Welter-Schultes (2012)

Family Trissexodontidae

Praeoestophorella phacodes (Thomä, 1845) 4G 107234, 107362 <1.0% RM† Yanes et al. (2009), Welter-Schultes (2012) [Caracollina Beck, 1837]

Table 1. Continued.


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give a full description of the specimens here, as this has already been done elsewhere; for this and a discussion of the main diagnostic features of each species, we refer the reader to the recent publications dealing with fossil

gastropods of MN5 age (e.g., Harzhauser et al. 2014, Salvador et al. 2015, 2016a, 2016b, Salvador & Rasser 2016). The best-preserved specimens found in the present material can be seen in Figures 2–4.

Figure 2. Fossil gastropods from Heuchlingen. For the inventory number of the material in the SMNS collection, please refer to Table 1. A–C. Pomatias conicus, 3 specimens, at different scales: (A) spire fragment, apertural view (H = 2.9 mm, D = 3.1 mm); (B) spire fragment, apical view (D = 3 mm); (C) operculum, external view (h = 5.1 mm, d = 4.4 mm). D. Radix socialis, juvenile, apertural view (H = 7.7 mm, D = 4.1 mm). E. Ferrissia deperdita (length = 4.8 mm). F–J. Gyraulus applanatus, 5 specimens, at the same scale: (F) H = 1.1 mm, D = 3.5 mm; (G) H = 0.9 mm, D = 3.5 mm; (H) H = 1.1 mm, D = 3.6 mm; (I) D = 3.4 mm; (J) D = 3.5 mm. K. Hippeutis subfontanus (H = 0.4 mm, D = 1.2 mm). L–M. Planorbarius mantelli, 2 specimens (at the same scale): (L) D = 6.6 mm; (M) D = 6.8 mm. N–P. Segmentina lartetii, 3 specimens (at the same scale): (N) H = 1.6 mm, D = 3.6 mm; (O) D = 3.6 mm; (P) D = 4.3 mm.


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Figure 3. Fossil gastropods from Heuchlingen (continued). For the inventory number of the material in the SMNS collection, please refer to Table 1. A. Deroceras sp. (H = 7.6 mm, D = 4.7 mm). B. Granaria sp. (H = 6.1 mm). C–E. Pseudidyla moersingensis, 3 fragments, not on the same scale: (C) spire apex fragment (H = 2.9 mm); (D) mid-spire fragment (H = 5.3 mm). (E) aperture fragment (H = 3.8 mm). F. Hypnophila loxostoma (H = 5.1 mm, D = 2.2 mm). G–I. Discus euglyphoides, 3 specimens (at the same scale): (G) apertural view (H = 1.9 mm, D = 3.8 mm); (H) apical view (D = 3.6 mm); (I) umbilical view (D = 4.2 mm). J–L. Discus pleuradrus, 3 specimens (at the same scale): (J) apertural view (H = 2.1 mm, D = 4.3 mm); (K) apical view (D = 3.5 mm); (L) umbilical view (D = 3.4 mm). M. Carychium eumicrum, at the same scale as C. galli (H = 1.4 mm, D = 0.6 mm). N. Carychium galli, at the same scale as C. eumicrum (H = 1.5 mm, D = 0.8 mm). O. Cecilioides aciculella (H = 2.0 mm, D = 0.8 mm). P. Gastrocopta acuminata, at the same scale as G. nouletiana (H = 2.5 mm, D = 1.5 mm). Q. Gastrocopta nouletiana, at the same scale as G. acuminata (H = 2.2 mm, D = 1.2 mm). R. Palaeotachea sp. (D = 10.9 mm). S. Pseudochloritis sp. (D = 7.3 mm). T, U. Protodrepanostoma involutum, 2 specimens (at the same scale): (T) apertural view (H = 2.2 mm, D = 3.8 mm); (U) apical view, spire collapsed into shell (D = 4.2 mm).


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Figure 4. Fossil gastropods from Heuchlingen (continued). For the inventory number of the material in the SMNS collection, please refer to Table 1. A–C. Leucochroopsis kleinii, 3 specimens (at the same scale): (A) apertural view (H = 4.1 mm, D = 5.6 mm); (B) apical view (D = 4.6 mm); (C) umbilical view (D = 4.4 mm). D. Urticicola perchtae (H = 2.3 mm, D = 4.6 mm). E. Opeas minutum (H = 4.4 mm, D = 1.8 mm). F. Oxyloma minima (H = 3.2 mm, D = 2.2 mm). G. Praeoestophorella phacodes (H = 1.8 mm, D = 4.0 mm). H. Negulopsis lineolata (H = 1.7 mm, D = 0.7 mm). I. Vertigo angulifera, at the same scale as V. callosa (H = 1.5 mm, D = 0.8 mm). J. Vertigo callosa, at the same scale as V. angulifera (H = 1.7 mm, D = 1.2 mm). K. Archaeozonites sp. (D = 7.8 mm).

In total, 30 gastropod species were found (Table 1), of which 6 are from freshwater habitats, while the remainder are terrestrial. All of these species are commonly found in coeval sediments from the Silvana Beds (e.g., Salvador et al. 2015, 2017, Salvador & Rasser 2016). By far the most abundant species are the freshwater planorbids Gyraulus applanatus and Planorbarius mantelli. Among the land snails, the most abundant are Gastrocopta spp., Opeas minutum, and Palaeotachea sp. Some species are represented by very few specimens (< 1% of relative abundance; Table 1) and occasionally just a single specimen was recovered (e.g., the fragile Cecilioides aciculella and Oxyloma minima).

From the species listed by Jooss in Kranz et al. (1924) for the region encompassing both Heuchlingen and Dettingen am Albuch (he did not differentiate where each specimen came from), the following species could not be found in the present material: Aegopinella subnitens (Klein, 1853), Apula coarctata (Klein, 1853), “Paludina” sp., Testacella zellii (Klein, 1853), and Triptychia kleini

Schnabel, 2006. The fragments of unionoid bivalves listed in Kranz et al. (1924) were likewise not found. The fragmentary Palaeotachea sp. and Pseudochloritis sp. in the present material (Table 1) refer probably to Palaeotachea renevieri (Maillard, 1892), Palaeotachea silvana (Klein, 1853), and Pseudochloritis incrassatus (Klein, 1853) listed by Kranz et al. (1924); however, poor preservation (only shell apexes are present) precluded their specific identification. Some of the listed species, however, were present in the voucher material of Jooss in the SMNS: Apula coarctata, Palaeotachea silvana, and Palaeotachea renevieri (Fig. 5). Finally, the pristine preservation of the vouchers of Vallonia subcyclophorella (Gottschick, 1911) indicates that they are actually contamination by Recent Vallonia costata (O.F. Müller, 1774).

PalaeoecologyFor the paleoecological analysis, the ecological prefer-ences of the Recent congeners of Heuchlingen’s fossils


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are listed in Table 1, alongside the references for such information. Although the Recent congeners of some spe-cies are known from too broad a range of habitats to have any use in a paleoenvironmental analysis, most are infor-mative. Furthermore, the ecological preferences of fossil genera have been treated in the paleontological literature and such interpretations are also used here (see Table 1 for references).

The Recent congeners of most of the freshwater spe-cies share a preference for richly vegetated, slow-moving, or standing waters (Table 1). Exceptions to this rule are: Segmentina, which is associated with marshy environ-ments, likely along the littoral area; and Ferrissia (and the hygrophilous land snail Oxyloma), which are typical inhabitants of reed-belts along the littoral.

The majority of land snails from Heuchlingen are deemed inhabitants of humid forests and/or scrublands (Table 1). Typical indicators of this kind of habitat are species with a hairy periostracum (e.g., Leucochro opsis, Protodrepanostoma, and Pseudochloritis), which is pre-served in fossil shells as tiny papillae (Fig. 4A). Further-more, Opeas and Negulopsis represent groups extinct in Europe that presently thrive in warmer climates. The genera Granaria and Praeoestophorella, however, are indicative of open calcareous meadows or rocky envi-ronments. Nevertheless, the relatively low abundance of these taxa could indicate that such habitats were rather removed from the depositional environment.

Of the species listed by Jooss in Kranz et al. (1924)

and not found within the present sample, some are worth-while mentioning. Jooss’s “Paludina” (Viviparidae) and his fragmentary unionoid bivalves would indicate the presence of deeper waters, on a more developed lake. Nevertheless, it is important to note that Jooss’s material come from limestones from other outcrops in the region, while the present material, given its preservation, is most likely from a marl. Two possibilities thus come to mind: (1) the 2 outcrops represent 2 consecutive stages of the lake, or (2) they represent 2 contemporaneous but distinct zones of the same lake (similar to what is known from the Silvana Beds of Randeck Maar; Rasser et al. 2013, Salvador et al. 2015).

ConclusionAccording to the rich freshwater and terrestrial gastropod paleocommunity of Heuchlingen, the locality’s paleo-environment can be summarized as a stagnant or slow moving permanent shallow water body surrounded by hu-mid woods with undergrowth and scrublands. Drier and likely calcareous meadows could be found further away.

AcknowledgementsWe are grateful to Karin Wolf-Schwenninger and Christina

Figure 5. Fossil gastropods from the region of Heuchlingen and/or Detting am Albuch found in the historical Jooss collection in the SMNS. A–C. Leucochroopsis kleinii (SMNS 66291; H = 5.9 mm, D = 7.9 mm). D. Palaeotachea renevieri, at the same scale as P. silvana (SMNS 66455; H = 11.0 mm, D = 12.7 mm). E. Palaeotachea silvana, on the same scale as P. renevieri (SMNS 66815; H = 17.9 mm, D = 19.7 mm). F–H. Planorbarius mantelli (SMNS 67863; H = 6.6 mm, D = 18.7 mm). I. Radix socialis (SMN S67693; H = 27.5 mm, D = 17.9 mm).


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G. Martin (SMNS) for the SEM images presented here, to Reinhard Ziegler and Elmar Heizmann (SMNS) for the data on the mammal remains and outcrop age, and to Dietrich Kadolsky, another (anonymous) reviewer, and the editor, John Hutchinson, for their helpful comments and suggestions. This work was partly supported by a Ph.D. grant from CNPq (Conselho Nacional de Desen-volvimento Científico e Tecnológico), Brazil, to R.B.S. (process no. 245575/2012-0).

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Manuscript submitted 13 March 2017 Revised manuscript accepted 23 September 2017

Documents

Fossil terrestrial and freshwater Gastropoda from the Eary ... · Rodrigo B. Salvador & Michael W. Rasser Staatliches Museum für Naturkunde Stuttgart, Rosenstein 1, 70191, Stuttgart,