Land and freshwater snails from the German Miocene

We studied several fossil outcrops in southern Germany, all dating from the Early to Middle Miocene. Back then, Germany was a subtropical forested land, sporting a huge diversity of snails. You can find a little more about the fossil sites and their molluscan fauna below.

Fossil sites

Below you can find a description of the paleoenvironment of each fossil site, i.e., how the environment was during the Miocene.

  • Adelschlag-Fasanerie: This carbonate-rich freshwater lake gradually became shallower throughout its duration.
  • Bechingen: Shallow temporary lakes or ponds surrounded by reeds and well-forested areas.
  • Biberach: The several localities from this region represent a system of meandering rivers and oxbow lakes.
  • Daugendorf: Shallow temporary lakes or ponds surrounded by reeds and well-forested areas.
  • Emerberg and Tautschbuch Hills: Varies. There are several localities along the southern margin of the Swabian Alb suitable for snails.
  • Gündlkofen: A small oxbow lake, remarkable by its almost complete lack of freshwater snails.
  • Heuchlingen: Shallow lake (or shallow margin of large lake) surrounded by humid forest and scrubland.
  • Hohenmemmingen: A humid and warm forest cover, with a well-vegetated and likely shallow water body.
  • Oggenhausen: A shallow lake or an oxbow lake, surrounded by lush vegetation.
  • Randeck Maar: The Randeck Maar is a volcanic crater (ca. 1.8 km diameter) which, after is formation during the Early Miocene, was filled by water and became a lake (with water depths of up to ca. 130 m).
  • Riedensheim: These were probably oxbow lakes or floodplain ditches in which the aquatic gastropods lived and the land snails were deposited at river highstands or during flash floods.
  • Sandelzhausen: It started as a small water-body, but gradually became a large lake. It has the most diverse fauna of all localities shown here.
  • Western Bavaria: The several fossil layers of these localities saw many changes, going from freshwater rivers and lakes, to brackish water environments and back to freshwater.

Snail diversity

Here you can see the number of mollusks species found on each fossil site. Further below, some of the more interesting species are highlighted.

  • Adelschlag-Fasanerie: 10 land snails + 7 freshwater;
  • Bechingen: 12 land snails + 7 freshwater;
  • Biberach, Ravensburg and Neu-Ulm: 10 land snails + 10 freshwater (including bivalves);
  • Daugendorf: 10 land snails + 3 freshwater;
  • Emerberg and Tautschbuch Hills: 50 land snails + 14 freshwater;
  • Gündlkofen: 18 land snails + 2 freshwater;
  • Heuchlingen: 24 land snails + 6 freshwater;
  • Hohenmemmingen: 18 land snails + 5 freshwater;
  • Oggenhausen: 12 land snails + 5 freshwater;
  • Randeck Maar: 28 land snails + 4 freshwater;
  • Riedensheim: 25 land snails + 9 freshwater;
  • Sandelzhausen: 30 land snails + 14 freshwater (including bivalves);
  • Western Bavaria: 6 land snails + 18 brackish/freshwater.

New species

The following new species were found and described:

  • Carychium (Carychium) galli Salvador, 2015 [Family Ellobiidae], from Sandelzhausen
  • Urticicola perchtae Salvador, 2013 [Family Hygromiidae], from Sandelzhausen



fig-localitiesA biogeography analysis was done to discover which land snail faunas (including most of those listed above and several others) were more similar to one another. We discovered that the snails from the two German states, Baden-Württemberg (marked as “BW” in the figure below) and Bavaria (“BY”), form two distinct groups, which are close to each other.



The preceding text is a summary of the following papers, where this research was published:

  • HÖLTKE, O.; SALVADOR, R.B.; RASSER, M.W. 2016. Paleobiogeography of Early/Middle Miocene terrestrial gastropods in Central Europe: an approach using similarity indices. Palaeogeography, Palaeoclimatology, Palaeoecology 461: 224-236.  [PDF]
  • HÖLTKE, O.; SALVADOR, R.B.; RASSER, M.W. 2018. Miocene continental gastropods from the southern margin of the Swabian Alb (Baden-Württemberg, SW Germany). Neues Jahrbuch fur Geologie und Palaontologie, Abhandlungen 287(1): 17–44.  [PDF]
  • SALVADOR, R.B. 2013. The fossil land and freshwater mollusks of Sandelzhausen (Early/Middle Miocene, Germany): Caenogastropoda, Neritimorpha, lower Heterobranchia and Bivalvia. Strombus 20: 19-26.  [PDF]
  • SALVADOR, R.B. 2013. The fossil pulmonate snails of Sandelzhausen (Early/Middle Miocene, Germany): Succineidae, Testacelloidea and Helicoidea. Zootaxa 3721: 157-171.  [PDF]
  • SALVADOR, R.B. 2014. The fossil land and freshwater snails of Gündlkofen (Middle Miocene, Germany). Zootaxa 3785: 271-287.  [PDF]
  • SALVADOR, R.B. 2015. The fossil pulmonate snails of Sandelzhausen (Early/Middle Miocene, Germany): Ellobiidae, Pupilloidea, and Clausilioidea. Palaontologische Zeitschrift 89: 37-50.  [PDF]
  • SALVADOR, R.B. & RASSER, M.W. 2014. The fossil pulmonate snails of Sandelzhausen (Early/Middle Miocene, Germany) (Hygrophila, Punctoidea and limacoids). Archiv fur Molluskenkunde 143: 187-202.  [PDF]
  • SALVADOR, R.B. & RASSER, M.W. 2016. Fossil land and freshwater gastropods from the Middle Miocene of Bechingen and Daugendorf, southwestern Germany. Archiv für Molluskenkunde 145(1): 111-124.  [PDF]
  • SALVADOR, R.B. & RASSER, M.W. 2016. The fossil land and freshwater snails of Oggenhausen (Middle Miocene, Germany). Revista Brasileira de Paleontologia 19(1): 41-52.  [PDF]
  • SALVADOR, R.B. & RASSER, M.W. 2017. Fossil terrestrial and freshwater Gastropoda from the Early/Middle Miocene of Heuchlingen, Germany. Archiv für Molluskenkunde 146(2): 233–241.  [PDF]
  • SALVADOR, R.B.; HÖLTKE, O.; RASSER, M.W. 2017. Fossil land and freshwater gastropods from the Miocene of Hohenmemmingen, Germany. Palaeodiversity 10: 41–48.  [PDF]
  • SALVADOR, R.B.; PIPPÈRR, M.; REICHENBACHER, B.; RASSER, M.W. Early Miocene continental gastropods from new localities of the Molasse Basin in southern Germany. Paläontologische Zeitschrift: 10.1007/s12542-016-0291-y  [PDF]
  • SALVADOR, R.B.; PRIETO, J.; MAYR, C.; RASSER, M.W. New gastropod assemblages from the Early/Middle Miocene of Riedensheim and Adelschlag-Fasanerie, southern Germany. Neues Jahrbuch fur Geologie und Palaontologie, Abhandlungen 279(2): 127-154.  [PDF]
  • SALVADOR, R.B.; RASSER, M.W.; HÖLTKE, O. 2015. Fossil gastropods from Miocene Lake Randeck Maar and its hinterland (SW Germany). Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, 277(3): 251-273.  [PDF]
  • SALVADOR, R.B.; SACH, V.J.; VALENTAS-ROMERA, B.L. 2015. The fossil continental mollusks in the Upper Freshwater Molasse (Middle Miocene) of the districts of Biberach, Ravensburg and Neu-Ulm, Germany. Revista Brasileira de Paleontologia 18(2): 201-216.  [PDF]