Paleobiogeography of the land snail communities of the Middle Miocene silvana-beds of Germany

Paleobiogeography of the land snail communities of the Middle Miocene silvana-beds of Germany | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Paleobiogeography of the land snail communities of the Middle Miocene silvana-beds of Germany Olaf Höltke, Rodrigo B. Salvador, Michael W. Rasser This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-2752635/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract The silvana -beds are the main component of the Upper Freshwater Molasse (“Obere Süßwassermolasse” or OSM) in southern Germany, Austria and Switzerland, named after the land snail Palaeotachea silvana (Klein, 1853 ). After initial disputes in the literature, in the 1920’s, malacologist Wilhelm Wenz better defined the silvana -beds lithology and its three informal subunits based on terrestrial gastropods. The silvana -beds mainly consist of limestone, marl and clay, eventually with sandy components. Despite remaining an informal stratigraphical unit, the silvana -beds is a practical tool for stratigraphic determination in the field and for paleomalacology studies. Information about the land snail communities of the silvana -beds are scattered in the literature and museum collections. Therefore, here we compile all this information and conduct a detailed biogeographic comparison of these localities’ snail faunas, based on similarity indices. For that purpose, all communities (containing at least six terrestrial gastropod species, from a total of 28 localities) within the silvana -beds and the adjoining coeval Bad Urach volcanic region were analyzed, based on four similarity indices and follow-up cluster analyses. The land snail communities of the silvana -beds formed six clusters (defined by ca. 50% similarity) according to the Sørensen-Dice index, five clusters by the Kulczynski and Ochiai indices, and four ones by the Simpson index. Despite some differences in the resulting dendrograms, enough similarity in the clustering was found for a meaningful comparison. The resulting clusters can be largely explained by geographic position, as is the case of the two main groupings recovered, representing the German Bavarian and Baden-Württemberg states. Some inconsistencies in the composition of the clusters might be explained by different stratigraphic positions within the silvana -beds, distinct paleoenvironmental conditions influencing the faunal composition, or the absence of key species. cluster analysis Gastropoda similarity indices Upper Freshwater Molasse Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 1. Introduction The silvana -beds (Middle Miocene) are the main component of the Upper Freshwater Molasse (“Obere Süßwassermolasse” or OSM) unit of the North Alpine Foreland Basin (NAFB) in southern Germany. A large quantity of often well-preserved continental mollusks, especially land snails, have been recovered from these beds, especially during the late 19th and early 20th centuries, being stored in several local and international museum collections. In spite of that, the silvana -beds have only rarely been subject to systematic excavations or intensive studies. After the pioneering works of Sandberger (1870–1875), Wenz (1923–1930) and others, thorough accounts of the silvana -beds molluscan faunas have only more recently started to appear, although still largely based on historical museum collections (Table 1). Our research group has been steadily collecting information on these fossil mollusks, revising and improving knowledge about them. As such, following the lines of Harzhauser & Mandic ( 2008 ) and Höltke et al. ( 2016 ), our objective here was to gather all information about the land snail communities of the many different outcrops of the silvana -beds and subject it to a paleobiogeographical analysis. In order to reach this goal, a thorough comparison of these snail faunas was conducted, using similarity indices and follow-up cluster analyses. 2. The Silvana-beds 2.1. Historical background Debates regarding the silvana -beds were exceedingly common in the geological literature of the 19th and early 20th centuries (good examples are Fraas 1882 and Rollier 1900 ). One of the main issues revolved around who first introduced this name and when. Simply enough, the silvana -beds are named after the land snail Palaeotachea silvana (Klein, 1853 ) (Schad 1908 ). According to Fraas ( 1888 ), the original material of P. silvana came from Mörsingen and Mundingen, but the label of the only syntype known names Zwiefalten (near Mörsingen) as type locality (Salvador et al. 2016a ). Despite being one of the most frequent fossil land snails in OSM deposits, in the species original description, Klein ( 1853 ) reported that it was fairly rare (“seltener”, in the original), so he did not mention any possibility of using it as an index fossil. However, the species identity was also up to debate, which caused confusion in the geological literature. Prior to Klein ( 1853 ) and for some time afterwards, the specimens from these beds were identified as Palaeotachea sylvestrina (Schlotheim, 1820 ) (e.g., Dunker 1848 ; Miller 1871 ), which is now understood to be a separate, younger species (Höltke & Rasser 2016 ). Fraas ( 1869 ) first proposed P. sylvestrina as an index fossil for the “Hohenmemmiger Kalke” (Hohenmemmingen’s limestone). Miller ( 1871 ) then used the term sylvestrina -beds for the OSM sediments in Hochsträß, near Ulm, without mention to P. silvana . Sandberger (1870–1875) then listed P. silvana from a several localities in the NAFB, but without considering its use as an index fossil. Quenstedt (1876), however, continued to use only the name P. sylvestrina as well as the term “Sylvestrinenkalke” ( sylvestrina -limestone). Deffner & Fraas ( 1877 ) then listed both species ( P. silvana and P. sylvestrina ) in their Tertiary chapter. Finally, Probst ( 1879 : 265–266) was the first to indicate P. silvana as an index fossil for the OSM; the identification of his material was done by Sandberger. Quenstedt ( 1884 ) still retained the name sylvestrina -beds, arguing that it was the common usage among geologists and that this species was known as P. silvana in the Swabian region. Fraas ( 1882 ) simply considered P. silvana as synonym of P. sylvestrina . However, Engel ( 1883 : 274) named the unit silvana -beds due to the index fossil P. silvana ; in his words, a very practical name (“sehr praktischer Namen”). Zittel ( 1885 ) then listed both species, P. silvana and P. sylvestrina , for the OSM, but without using the terms silvana - or sylvestrina -beds. Fraas ( 1888 ) finally adopted the term silvana -limestone (“Silvanakalk”), but still considered P. silvana and P. sylvestrina to represent the same species. In summary, from the first use of P. silvana as index fossil (Probst, 1879 ), through the naming of the unit as silvana -beds (Engel, 1883 ) and onwards, several different opinions remained in the literature. These concerned both the unit’s name and the validity of P. silvana as a separate species. Nevertheless, by the beginning of the 20th century, the name silvana -beds prevailed (e.g., Rollier 1900 , 1903 ; Miller 1901 , 1903 ; Jooss 1915 ; Gottschick & Wenz 1916 ). Still, while some authors started to focus on describing lithological subunits within the silvana -beds (Schad 1908 for the Ehingen/Donau region; Schwarz 1913 , for the Emerberg/Tautschbuch region), another issues arose: the stratigraphical position and age of the beds. Researchers considered two main possibilities, either Upper Oligocene (Rollier 1900 ) or Upper Miocene (Miller 1901 ; Jooss 1915 ; Dietrich & Kautsky, 1920 ). Finally, Wenz ( 1920 ) summarized the then current knowledge and published a classification of the silvana beds, dividing them in three sections based on the land snail fauna (Fig. 3 ; see also Höltke & Rasser 2016 ) and placing it in the Upper Miocene (Tortonian). Furthermore, Wenz ( 1921 ) separated the units into the older silvana -beds and the younger sylvestrina -beds. The lowest section of the silvana -beds received a special name, “Flammenmergel” or “flame marl” (Wenz 1924 ). In the decades that followed, publications about the silvana -beds became much scarcer, with some accounts of fossil snails (Gottschick & Wenz 1927 ; Schlickum 1976 ) and regional geological works (Haag 1960 ; Zöbelein 1977 ). Only much later, the silvana -beds started to draw renewed attention (Schweigert 1996 ; Esu 1999 ; Doppler et al. 2005 ). Esu ( 1999 ) was responsible for placing the silvana -beds within the Neogene Mammal zone MN5 of the Middle Miocene); hence, these beds predate the Nördlinger Ries and Steinheim am Albuch meteorite events. Starting with Sach ( 1999 ), the molluscan fauna of the silvana -beds began to be re-examined (see publications listed in Table 1). Finally, Höltke & Rasser ( 2016 ) revised the Palaeotachea species from the German Miocene and discussed the implications for stratigraphy. 2.2. Stratigraphy and lithology As explained above, the silvana -beds are a unit within the OSM in the North Alpine Foreland Basin (for details about this basin, see Kälin & Kempf 2009 ), defined by the presence of the index fossil Palaeotachea silvana , a terrestrial pulmonate gastropod. The strata underlying the silvana -beds are region-dependant and can be: the “Helicidenmergel” (“helicid-marl”), the Upper Brackish Water Molasse (“Brackwassermolasse”) or the Upper Jusassic Malm. Immediately above the silvana -beds are the silvestrina -beds (Höltke & Rasser 2016 ) or, where the latter is absent, Pleistocene gravel. The silvana -beds are freshwater sediments, consisting of limestone, marl, silt and also partly with a sandy component. Naturally, they also include several fossils, especially gastropods. In particular, the deposits of the Emerberg/Tauschbuch region at the southern margin of the Swabian Alb (Fig. 1 ) are a hotspot of diversity and preservation of land snails (Höltke et al. 2018 ). Wenz ( 1920 , 1921 , 1924 ) divided the silvana -beds in three different subunits based mainly on their land snail fauna (Fig. 2 ). Despite some criticisms (Jooss 1923 ; Zöbelein 1977 ) regarding wider stratigraphic distribution of some species, the structuring of the silvana -beds sensu Wenz is a practical tool, especially when dealing with the usually incomplete information available for most localities. Furthermore, more region-specific descriptions and subdivisions of the silvana -beds were published by Schad ( 1908 ), Schwarz ( 1913 ) (see Fig. 3 ), Haag ( 1960 ) and Zöbelein ( 1977 ); the latter also provided a critical overview about the works of the former three authors. 3. Material And Methods We compiled literature data on all localities belonging to the silvana -beds, using the most up to date works available. Moreover, the coeval localities from the neighboring Bad Urach volcanic region of the Swabian Alb (which lies just outside the NAFB) were included. As a compromise between the scarcity of data for several localities and a meaningful number for the analyses using similarity indices, we only considered those localities with at least six species of land snails. The chosen localities are listed in Table 1. In some cases, the literature available dates from the early 20th century, so we corrected the species lists they presented regarding minor taxonomic issues (synonymies and new names) according to the more recent literature (Table 1). In a few cases, literature data could be complemented with material from museum collections, namely: Staatliches Museum für Naturkunde Stuttgart (SMNS; Stuttgart, Germany), Bayerische Staatssammlung für Paläontologie und Geologie (BSPG; Munich, Germany), and Department of Geosciences of the Eberhard Karls Universität Tübingen (GPIT; Tübingen, Germany; formerly Geologisch-Paläontologisches Institut Tübingen). The resulting list of land snail species for each locality was then subjected to a cluster analysis using four distinct similarity indices with “Paired group (UPGMA)” algorithm. The indices used for the presence-absence data of the species were: Sørensen-Dice (Dice 1945 ; Sørensen 1948 ), Kulczynski ( 1927 ), Ochiai ( 1957 ), and Simpson ( 1943 , 1960 ). This was done to test for stability and consistency of the groupings in the resulting dendrograms. In this step, we removed those species endemic to a single site. The analyses were conducted with 10,000 bootstrap replicates to test for the robustness of the groups (robustness estimates, in percentages, are shown at each node of the dendograms). The analyses were carried out in PAST v.3.16 (Hammer et al. 2001 ). Finally, we searched for possible common land snail micro-communities, that is, a constant assemblage of some species that could be found on more than one locality. However, the cluster analysis results were inconsistent, possibly due to the small sample size of most localities or slight differences in stratigraphical position between geographically close localities. Therefore, this step is omitted in the present article. 4. Results A total of 28 localities were used in the present work (Table 1), containing together 51 different land snail species. The localities with the highest number of species are Mörsingen and Zwiefaltendorf, with 36 each. The dendrograms resulting from the cluster analyses (Figs. 4 –7) share similarities and differences. To conduct comparisons between them, only clusters with a similarity index of ca. 50% were taken into account (for the Simpson index, subgroups with ca. 65% similarity were explored; Fig. 6 ). The clusters were numbered in Figs. 3 – 6 and these numbers are used throughout the text. The robustness estimates from the bootstrap are usually low for the large clusters, but can achieve high values for some locality pairs (e.g., Mörsingen and Zwiefaltendorf). Low values normally indicate that the cluster is not well supported; however, in the present case the low values might be exacerbated due to the small sample size of several terminals (localities) and the thus disproportionate importance of the few species-rich terminals inside the clusters. Robustness estimates values are especially low for the Simpson index, but the results of this index are also carefully discussed herein as it is deemed very appropriate for paleontological samples in which there are large differences in species richness between localities (Simpson 1943 , 1960 ). The Bavarian localities Riedensheim, Sandelzhausen and Adelschlag-Fasanerie form one reasonably well-supported cluster in all indices except for Simpson, which includes other localities (Figs. 4 –7: clusters D1, K1, O1, S1). In all dendrograms, there is one large cluster containing most of the Baden-Württemberg deposits located at the Southern and Eastern margin of the Swabian Alb (Figs. 4 –7: clusters D2, K2, O3, S2). In all cases, the Bavarian locality Gündlkofen is clustered with the Austrian Pfänder region (Figs. 4 –7: clusters D6, K5, O6, S4). The localities Georgensgmünd and Mundingen form one cluster by the Sørensen-Dice and Kulczynski indices, but include Ehingen/Donau by the Ochiai index (Figs. 4 – 6 : clusters D5, K4, O4); this cluster is absent in the Simpson index dendrogram. In all dendrograms, the localities Dächingen, Randecker Maar, Hengen and Oggenhausen are clustered together (Figs. 4 –7: clusters D3, K3, O2, S3), but by the Ochiai index, it also contains the localities Daugendorf and Andelfingen. There is an additional large cluster identified by the Sørensen-Dice index (D4), containing four localities that are scattered through other clusters according to the other indices. The locality Baach is separated from all others in all four indices and the position of the locality Vermes is extremely variable. In the dendrogram resulting from Simpson index (Fig. 7), there are five pairs of localities with 100% similarity. This is because this index is especially good in detecting nestedness of samples, that is, if one community contains the same species as another community and a few extra ones, the latter is nested within the former (index value equals one). 5. Discussion The Dice, Kuclczynski and the Ochiai index plots are overall very similar to one another, whereas the Simpson index plot is the most different. Nevertheless, despite these differences, the basic arrangement is still similar enough for meaningful comparison. The similarities of clusters are discussed in further detail below and possible reasons for the observed differences are offered. 5.1. Bavarian cluster The cluster containing the Bavarian localities Riedensheim, Sandelzhausen and Adelschlag-Fasanerie (Figs. 4 –7: clusters D1, K1, O1, S1) can be explained by their geographic position: the three deposits are geographically very close. These localities have also been deemed very similar in previous works (Höltke et al. 2016 ; Salvador et al. 2016b ), which have also included Undorf, another closely located deposit. Undorf, however, here is included in the Baden-Württemberg cluster (see below) for all indices except Simpson (Fig. 7). By the latter, Undorf is featured together with the three localities above and also the Swiss deposits of Vermes. 5.2. Vermes The only Swiss locality, Vermes, has a very variable position in the dendrograms. The locality is generally well separated from the others, with low similarity values. According to the Sørensen-Dice and Ochiai indices (Figs. 4 , 6 ), it is more closely related to Baden-Württemberg localities, while the Kulczynski and Simpson indices place it closer to the Bavarian localities. In fact, by the latter index, it is placed well inside the Bavarian group (see above), together with the locality Undorf. The reason for this clustering, despite their geographic distance, could be the high number of species it shares with Undorf (seven: Archaeozonites costatus , Gastrocopta nouletiana , Leucochroopsis kleini , Palaeotachea silvana , Testacella zelli , Triptychia kleini , and Vitrina suevica ). 5.3. Baden-Württemberg cluster This large cluster can be found, with a few differences, in all indices (Figs. 4 –7: clusters D2, K2, O3, S2). Most of the localities are located on the southern margin of the Swabian Alb (Fig. 1 ), which is a likely reason behind their similarity in species composition. Dischingen and Hohenmemmingen, in particular, are slightly removed towards the eastern margin of the Alb, but still fairly close. An unusual locality within this cluster is Undorf, located in Bavaria. Undorf is often deemed more related to the Bavarian cluster (Höltke et al. 2016 ; Salvador et al. 2016b ), which is more reasonable from a paleogeographical point of view. In fact, the results of the Simpson index places Undorf within the Bavaria cluster (Fig. 7). The reason for the grouping of Undorf with the Baden-Württemberg localities might be due to the 21 species it shares with the joint localities from the southern margin of the Swabian Alb, because the other indices used herein give more weight to shared species between localities. Other inconsistencies are the presence of the Bavarian locality Georgensgmünd by Simpson index (Fig. 7) and the absence of Friedingen (near to Zwiefalten and thus withing the geographical region) according to Sørensen-Dice index. The localities within this cluster may represent the lower and middle silvana -beds ( sensu Wenz 1920 , 1921 , 1924 ) (see also Fig. 3 ) due to the abundance of the species P. silvana and Pseudochloritis incrassata , as well as the occurrence of Palaeotachea dentula in some deposits. 5.4. Maar cluster This cluster is made up of “geographically mixed” localities (Figs. 4 –7: clusters D3, K3, O2, S3), with most localities “scattered” through Baden-Württemberg state. According to Sørensen-Dice and Simpson indices, this clusters consist of Randecker Maar, Hengen, Dächingen and Oggenhausen (Figs. 4 , 7: clusters D3, S3), which share overall five species ( Archaeozonites costatus , Palaeotachea renevieri , P. silvana , Pomatias conica , and Pseudochloritis incrassata ). The Kulzynski index also includes Andelfingen (Fig. 5 : cluster K3), resulting in overall three species in common ( A. costatus, P. silvana, P. incrassata ). Finally, the Ochiai index includes both Andelfingen and Daugendorf (Fig. 6 : cluster O2), resulting in only two species in common (P. silvana, P. incrassata ). The pairing of the localities Randeck Maar and Hengen in all indices except Simpson, could be related to the fact that both had a maar lake paleoenvironment. Dächingen and Oggenhausen had very different paleoenvironments, but the few species shared with the others may have influenced this clustering. The reasons for the inclusion of Daugendorf and Andelfingen are unclear. They have only the two most typical species in common with the other four localities ( P. silvana & P. incrassata ). According to the presence of P. incrassata the reason for this clustering maybe the same stratigraphic position of deposits named above (Middle silvana -beds sensu Wenz 1924 ) (see also Fig. 3 ). 5.5. Cluster D4 The cluster D4 appears exclusively on the dendrogram from the Sørensen-Dice index (Fig. 4 ). It only contains deposits located on the southern margin of the Swabian Alb, but not included in the Baden-Würtemberg cluster (see above) by this index. In all other indices, the localities forming the D4 cluster are usually part of the Baden-Würtemberg cluster. All the localities have only three species in common: Leucochroopsis kleini , Palaeotachea silvana , Pseudochloritis incrassata . The reasons for the Sørensen-Dice index separating this second Swabian Alb cluster are unclear, but may be related to the absence of some key species or to different stratigraphic positions. 5.6. Georgensgmünd/Mundingen clusters In all indices except for Simpson, Mundingen (part of the Ehingen/Donau district) in Baden-Württemberg forms one cluster with the Bavarian locality Georgensgmünd (Figs. 4 – 6 : clusters D5, K4 and O4). Despite their different geographic positions, the two localities have four species in common ( Apula coarctata, Janulus supracostatus , P. silvana , and Pseudochloritis incrassata ). The Ochiai index also includes the locality Ehingen/Donau, which is close to Mundingen. Nevertheless, the three deposits only share the two typical silvana -beds species ( P. silvana and Pseudochloritis incrassata ). The reason for this clustering is also unclear, but could be related to similar stratigraphic positions within the silvana -beds or to the influence of paleoenvironmental factors. 5.7. Gündlkofen/Pfänder cluster The Bavarian locality Gündlkofen and the Austrian deposit Pfänder (near Bregenz) form one cluster, quite isolated from all others (Figs. 3 – 6 : D6, K5, O5, S4). The two localities have four species in common: P. silvana, Pseudidyla moersingensis, Pseudochloritis incrassata, Triptychia teutonica . The reasons for the clustering might be either geographic or stratigraphic, as Gündlkofen is deemed to be younger than the other Bavarian localities. 5.8. Baach The locality Baach (part of the Zwiefalten district) is located on the southern margin of the Swabian Alb in Baden-Württemberg, but it is completely isolated from all other localities in all four cases. All the species occurring in Baach can also found in the deposits from the Swabian Alb. The most important features of Baach are the missing of of Pseudochloritis incrassata and the presence of Palaeotachea dentula . So the reason for the separation may be due to the stratigraphic position representing the lower silvana -beds sensu Wenz 1924 (see also Fig. 3 ). 6. Conclusion The different clusters in the present biogeographic analyses can be largely explained by their geographic position, like the Bavarian and Baden-Württemberg clusters. Some inconsistencies in the composition of the clusters might be explained by different stratigraphic positions within the silvana -beds (Georgensgmünd/Mundingen and Gündlkofen/Pfänder clusters), distinct paleoenvironmental conditions influencing the faunal composition (Maar cluster) or absence of key species (maybe Baach). However, with the scarce data presently available, it is difficult to consider the likelihood of such hypotheses. At present, the silvana -beds (and also the sylvestrina -beds) have not been defined according to international stratigraphical guidelines, thus remaining an informal lithostratigraphic unit. The only distinct feature of the silvana -beds is the presence of the land snail P. silvana , which is also true for the sylvestrina -beds and P. sylvestrina . Perhaps it would be sensible to define it as biostratigraphic zone, in the lines of the Mesozoic ammonoid zones. In this way, the coeval sediments from the nearby Bad Urach volcano region (e.g., Hengen), which is just outside the NAFB and bear the same land snail fauna, could be included without problems in the silvana -beds. However, for detailed stratigraphic research on the silvana -beds, good outcrops/profiles would be necessary, which are unfortunately unavailable in the present. Declarations Funding: None. Conflicts of interest/Competing interests: There are no conflicts of interest. 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Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, 287(1): 17–44. http://dx.doi.org/10.1127/njgpa/2018/0704 Jooss, C.H. (1910). Binnenconchylienfauna aus dem Obermiozän des Pfänders bei Bregenz am Bodensee. Nachrichtsblatt der Deutschen Malakozoologischen Gesellschaft , 42 : 19– 29. Jooss, C.H. (1915). Zur Altersfrage der Süßwasserablagerungen bei der Ruggburg am Pfänder bei Bregenz. Centralblatt für Mineralogie, Geologie und Palaeontologie , 1915: 62–64. Jooss, C.H. (1923). Die Schneckenfauna der süddeutsch-schweizerischen Helicidenmergel und ihre Bedeutung für die Altersbestimmung der letzteren. Neues Jahrbuch für Geologie und Paläontologie , 49: 185 ̶ 211. Kälin, D., & Kempf, O. (2009). High-resolution stratigraphy from the continental record of the Middle Miocene Northern Alpine Foreland Basin of Switzerland. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen , 254: 177–235. http://dx.doi.org/10.1127/0077-7749/2009/0010 Klein, A. von (1846). Conchylien der Süsswasserkalkformation Württembergs. J ahreshefte des Vereins für vaterländische Naturkunde in Württemberg , 2: 60–116. Klein, A. von (1853). Conchylien der Süßwasserkalkformation Württembergs. Jahreshefte des Vereins für vaterländische Naturkunde in Württemberg , 9: 203–223. Kulczynski, S. (1927). Die Pflanzenassoziationen der Pieninen. Bulletin International de l’Academie Polonaise des Sciences et des Lettres, Classe des Sciences Mathematiques et Naturelles B , 1927: 57–203. Maassen, W.J.M. (2003). Notes on terrestrial molluscs of the island of Sulawesi. 3. The genera Palaina, Arinia and Opisthostoma (Gastropoda, Prosobranchia, Diplommatinidae), with descriptions of a dozen new taxa. Basteria , 67: 47– 63. Miller, K. (1871). Das Tertiär am Hochsträss. Jahreshefte des Vereins für vaterländische Naturkunde in Württemberg, 27: 272–292. Miller, K. (1901). Zum Alter des Sylvanakalkes. Centralblatt für Mineralogie, Geologie und Palaeontologie , 1901: 129–133. Miller, K. (1903). Zu Rollier, das Alter des Sylvanakalks.- Centralblatt für Mineralogie, Geologie und Palaeontologie , 1903: 141–144. Nekola, J.C. & Coles, B.F. (2010). Pupillid Land Snails of Eastern North America. American Malacological Bulletin , 28(2): 29–57. http://dx.doi.org/10.4003/006.028.0221 Ochiai, A. (1957). Zoographic studies on the soleoid fishes found in Japan and its neighboring regions. Bulletin of the Japanese Society of Scientific Fisheries , 22: 526–530. Probst, J. (1879). Verzeichnis der Fauna und Flora der Molasse im württembergischen Oberschwaben. Jahreshefte des Vereins für vaterländische Naturkunde in Württemberg , 35: 221–304. Quenstedt, F.A. (1884). Begleitworte zur geognostischen Specialkarte von Württemberg. Atlasblätter Ehingen, Biberach, Laupheim, Ochsenhausen . A. Kleeblatt & Comp. Quenstedt, F.A. (1884). Petrefactenkunde Deutschlands. 7. Band Gasteropoden . 867 pp. Leipzig (Fue`s). Rasser, M.W., Bechly, G., Böttcher, R., Ebner, M., Heizmann, E.P.J., Höltke, O., Joachim, C., Kern, A.K., Kovar-Eder, J., Nebelsick, J.H., Roth-Nebelsick, A., Schweigert, G., Schoch, R.R., Ziegler, R. (2013). The Randeck Maar: Palaeoenvironment and habitat differentiation of a Miocene lacustrine system. Palaeogeography, Palaeoclimatology, Palaeoecology, 392: 426– 453. http://dx.doi.org/10.1016/j.palaeo.2013.09.025 Rollier, L. (1900). Vorläufige Notiz über das Alter des Sylvanakalkes. Centralblatt für Mineralogie, Geologie und Palaeontologie , 1900: 89–91. Rollier, L. (1903). Über Diskordanzen im Schwäbischen Tertiär. Vierteljahrsschrift der Naturforschenden Gesellschaft Zürich , 48: 307–320. Sach, J. (1999). Litho- und biostratigraphische Untersuchungen in der Oberen Süßwassermolasse des Landkreises Biberach a. d. Riß (Oberschwaben).- Stuttgarter Beiträge zur Naturkunde , Serie B, 276:1-167. Salvador, R.B. (2013a). The fossil land and freshwater mollusks of Sandelzhausen (Early/Middle Miocene, Germany): Caenogastropoda, Neritimorpha, lower Heterobranchia and Bivalvia. Strombus , 20(1-2): 19–26. Salvador, R.B. (2013b). The fossil pulmonate snails of Sandelzhausen (Early/Middle Miocene, Germany): Succineidae, Testacelloidea and Helicoidea. Zootaxa , 3721(2): 157–171. http://dx.doi.org/10.11646/zootaxa.3721.2.3 Salvador, R.B. (2014). The fossil land and freshwater snails of Gündlkofen (Middle Miocene, Germany). Zootaxa , 3785(2): 271–287. https://doi.org/10.11646/zootaxa.3785.2.9 Salvador, R.B. (2015). The fossil pulmonate snails of Sandelzhausen (Early/Middle Miocene, Germany): Ellobiidae, Pupilloidea, and Clausilioidea. Paläontologische Zeitschrift , 89(1): 37–50. http://dx.doi.org/10.1007/s12542-013-0210-4 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. http://dx.doi.org/10.1127/arch.moll/1869-0963/143/187-202 Salvador, R.B. & Rasser, M.W. (2016a). Fossil gastropods from the Middle Miocene of Bechingen and Daugendorf, southwestern Germany. Archiv fur Molluskenkunde , 145(1): 111– 124. DOI: 10.1127/arch.moll/1869-0963/145/111-124 Salvador, R.B. & Rasser, M.W. (2016b). The fossil land and freshwater snails of Oggenhausen (Middle Miocene, Germany). Revista Brasileira de Paleontologia , 19(1): 41–52. http://dx.doi.org/10.4072/rbp.2016.1.04 Salvador, R.B., Höltke, O. & Rasser, M.W. (2017). Fossil land and freshwater gastropods from the Miocene of Hohenmemmingen, Germany. Palaeodiversity , 10: 41–48. http://dx.doi.org/10.18476/pale.v10.a4 Salvador, R.B. & Höltke, O. & Rasser, M.W. (2018). Miocene continental gastropods from Dischingen, Germany. Palaeodiversity , 11(1):11–19. http://dx.doi.org/10.18476/pale.11.a2 Salvador, R.B., Höltke, O., Rasser, M.W. & Kadolsky, D. (2016a). Annotated type catalogue of the continental fossil gastropods in the Staatliches Museum für Naturkunde Stuttgart, Germany. Palaeodiversity , 9: 15–70. http://dx.doi.org/10.18476/pale.v9.a3 Salvador, R.B., Prieto, J., Mayr, C., Rasser, M.W. (2016b). New gastropod assemblages from the Early/Middle Miocene of Riedensheim and Adelschlag-Fasanerie, southern Germany. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen , 279: 127–154. http://dx.doi.org/10.1127/njgpa/2016/0546 Salvador, R.B., Rasser, M.W. & Höltke, O. (2015a). Fossil gastropods from Miocene Lake Randeck Maar and its hinterland (SW Germany). Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen , 277(3): 251–273. http://dx.doi.org/10.1127/njgpa/2015/0505 Salvador, R.B, Sach, V. & Valentas-Romera, B.L. (2015b). 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. http://dx.doi.org/10.4072/rbp.2015.2.02 Sandberger, F. (1870–1875). Die Land- und Süßwasserconchylien der Vorwelt : livr. 1: 1– 48, pl. 1– 4, 1870; livr. 2– 3: 49– 96, pl. 5– 12, 1870; livr. 4– 5: 97– 160, pl. 13– 20, 1871; livr. 6– 8: 161– 256, pl. 21– 32, 1872; livr. 9– 10: 257– 352, pl. 33– 36, 1873; livr. 11: 353– 616, 1875; livr. 12: 617– 1000, 1875. Kreidel. Schad, J. (1908). Beitrag zur Kenntnis des Tertiärs am Landgericht und Hochsträß. Jahreshefte des Vereins für vaterländische Naturkunde in Württemberg , 64: 249–304. Schlickum, W.R. (1976). Die in der pleistozänen Gemeindekiesgrube von Zwiefaltendorf a. d. Donau abgelagerte Molluskenfauna der Silvanaschichten. Archiv für Molluskenkunde 107(1/3): 1–31. Schlotheim, E.F. (1820). Die Petrefaktenkunde auf ihrem jetzigen Standpunkt durch die Beschreibung seiner Sammlung versteinerter und fossiler Überreste des Thier- und Pflanzenreichs der Vorwelt erläutertBecker-Verlag. Schwarz, F. (1913). Beschreibung des Tertiärs im Tautschbuch-Emerberggebiet . 54 pp. Inaugural-Dissertation, Tübingen (Laupp). Schweigert, G. (1996). Vergleichende Faziesanalyse, Paläoökologie und palöogographisches Umfeld tertiärer Süßwasserkarbonate auf der westlichen Schwäbischen Alb und im Hegau (Baden-Württemberg). Profil , 9: 1–100. Simpson, G.G. (1943). Mammals and the nature of continents. American Journal of Science , 241: 1–31. http://dx.doi.org/10.2475/ajs.241.1.1 Simpson, G.G. (1960). Notes on the measurement of faunal resemblance. American Journal of Science , 258A: 300–311. Sørensen, T. (1948). A method of establishing groups of equal amplitude in plant sociology based on similarity of species and its application to analyses of the vegetation on Danish commons. Kongelige Danske Videnskabernes Selskab , 5(4): 1–34. Wedel, J. (2008). Pleistocene molluscs from research boreholes in the Heidelberg Basin. Quaternary Science Journal , 57(3–4): 382–402. http://dx.doi.org/10.3285/eg.57.3-4.6 Wenz, W. (1920). Über das Vorkommen von Cepaea eversa larteti (Boissy) in den schwäbischen Silvanaschichten und seine Bedeutung für deren Gliederung. Senckenbergiana, 2:151–158. Wenz, W. (1921). Zur Frage der Altersstellung des schwäbischen Tertiärs. Centralblatt für Mineralogie, Geologie und Palaeontologie, 1921:559–564. Wenz, W. (1923-1930). Gastropoda Extramarina Tertiaria. I-X . W. Junk. Wenz, W. (1924). Die Flammenmergel der Silvanaschichten und ihre Fauna. Jahresberichte und Mitteilungen des Oberrheinischen geologischen Vereins , N.F., 13: 181–186. Wenz, W. (1933). Zur Land- und Süßwassermolluskenfauna der subalpinen Molasse des Pfändergebietes. Senckenbergiana , 15: 7–12. Wenz, W. (1935). Weitere Beiträge zur Land- und Süßwasser-Molluskenfauna der subalpinen Molasse des Pfändergebietes. Senckenbergiana , 17: 223–226. Werner, W. (2014). Gauinger, Sonderbucher und Riedlinger Travertin . In: Werner, W., Wittenbrink, J., Bock, H. & Kimmig, B. (Eds.), Naturwerksteine aus Baden-Württemberg. Vorkommen, Beschaffenheit und Nutzung (pp. 279–294). Landesamt für Geologie, Rohstoffe und Bergbau. Zittel, K.A. (1885). Handbuch der Palaeontologie. 1. Abteilung Palaeozoologie. II Band. Mollussca und Arthropoda . Oldenbourg. Zöbelein, H.K. (1977). Anstehende und eiszeitlich verschleppte Obere Süßwasser-Molasse (Ober-Miozän) im Bereich des Riedlinger Beckens (Baden-Württemberg). Mitteilungen der Bayerischen Staatssammlung für Paläontologie und Historische Geologie , 17: 291–334. Table 1 Table 1. Deposits and references. Locality Inside NAFB? Country Nr. Species Reference(s) Adelschlag-Fasanerie yes Germany (BY) 8 Salvador et al. (2016b) Altheim near Ehingen/Donau yes Germany (BW) 13 Höltke et al. (2018) Andelfingen yes Germany (BW) 8 Höltke et al. (2018) Baach yes Germany (BW) 7 Höltke et al. (2018) Bechingen yes Germany (BW) 17 Salvador & Rasser (2016a) Dächingen yes Germany (BW) 6 Höltke et al. (2018) Daugendorf yes Germany (BW) 7 Salvador & Rasser (2016) Dischingen yes Germany (BW) 25 Salvador et al. 2018 Ehingen/Donau yes Germany (BW) 6 Höltke et al. (2018) Emerberg yes Germany (BW) 18 Höltke et al. (2018) Friedingen yes Germany (BW) 8 Wenz (1923) Georgensgmünd yes Germany (BY) 7 Berger (2010) Gündlkofen yes Germany (BY) 7 Salvador (2014) Hausen ob Allmendingen yes Germany (BW) 11 Höltke et al. (2018) Hengen no Germany (BW) 11 Ehrat & Jooss (1921), collection material Hohenmemmingen yes Germany (BW) 24 Salvador et al. (2017) Mörsingen yes Germany (BW) 36 Schlickum (1976), collection material Mundingen yes Germany (BW) 6 Wenz (1923) Oggenhausen yes Germany (BY) 8 Salvador & Rasser (2016b) Pfänder near Bregenz yes Austria (Vorarlberg) 8 Jooss (1910, 1915), Wenz (1933, 1935) Randecker Maar no Germany (BW) 17 Salvador et al. (2015a) Rennertshofen-Riedensheim yes Germany 13 Salvador et al. (2016b) Sandelzhausen yes Germany (BW) 14 Salvador (2013a, 2013b, 2015); Salvador & Rasser (2014) Stoffelberg near Ehingen/Donau yes Germany (BW) 14 Höltke et al. (2018) Undorf yes Germany (BY) 28 Clessin (1911) Vermes yes Switzerland (Jura) 8 Wenz (1923) Zwiefalten yes Germany (BW) 17 Höltke et al. (2018) Zwiefaltendorf yes Germany (BW) 36 Schlickum (1976), Höltke et al. (2016) Cite Share Download PDF Status: Posted Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-2752635","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":193644182,"identity":"5b1fa81e-9215-411d-bf0b-d56a4b7f9302","order_by":0,"name":"Olaf Höltke","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAAvklEQVRIiWNgGAWjYDACdgY2IGkhx8DAA+IyE6GFGaxFwph0LYkNRGvhB2p58KFCIn3D8bMHH3xgsJYjqEWymYHdcMYZidwNZ/KSDWcwpBsT1GJwmIFNmrcNqOVAjpk0D8NhoAuJ0vJPIt3g/BuwlnoitTRIJBjcgNiSQIxf2CRnHJMwnHnjjbHhDIN0Q4K28LM3sEl8qLGR5zufYwgMOmt5grYANX0AUwoHwO4kQgMcyBN00CgYBaNgFIxYAAB0pjJnw7F3NAAAAABJRU5ErkJggg==","orcid":"","institution":"Staatliches Museum für Naturkunde Stuttgart: Staatliches Museum fur Naturkunde Stuttgart","correspondingAuthor":true,"submittingAuthor":false,"prefix":"","firstName":"Olaf","middleName":"","lastName":"Höltke","suffix":""},{"id":193644183,"identity":"fc0215e0-162d-43eb-8b0d-f9c0985dc739","order_by":1,"name":"Rodrigo B. Salvador","email":"","orcid":"","institution":"Department of Arctic and Marine Biology at the University of Tromso, Norway","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Rodrigo","middleName":"B.","lastName":"Salvador","suffix":""},{"id":193644184,"identity":"fa49de5d-3870-4a1a-976a-eb3973985c07","order_by":2,"name":"Michael W. Rasser","email":"","orcid":"","institution":"Staatliches Museum für Naturkunde Stuttgart: Staatliches Museum fur Naturkunde Stuttgart","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Michael","middleName":"W.","lastName":"Rasser","suffix":""}],"badges":[],"createdAt":"2023-03-29 15:13:11","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-2752635/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-2752635/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":36275642,"identity":"546046b3-61dc-4bc5-b6d3-7d4be8096b8e","added_by":"auto","created_at":"2023-04-25 13:25:34","extension":"jpg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":685664,"visible":true,"origin":"","legend":"\u003cp\u003eGeographic position of the single localities.\u003c/p\u003e","description":"","filename":"Fig.1.jpg","url":"https://assets-eu.researchsquare.com/files/rs-2752635/v1/bfc5a61b300e19cf356cf919.jpg"},{"id":36275134,"identity":"9dba5c9f-3315-4a3f-b6bb-8aabca345469","added_by":"auto","created_at":"2023-04-25 13:17:34","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":618437,"visible":true,"origin":"","legend":"\u003cp\u003eStructuring of the \u003cem\u003esilvana\u003c/em\u003e-beds (redrawn from Wenz (1924) and Höltke \u0026amp; Rasser (2016)\u003c/p\u003e","description":"","filename":"Fig.2.jpg","url":"https://assets-eu.researchsquare.com/files/rs-2752635/v1/52c1f1d13eed976a615a7169.jpg"},{"id":36274227,"identity":"4a3ffc7e-531a-4ca2-8482-62143908efbb","added_by":"auto","created_at":"2023-04-25 13:09:34","extension":"jpg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":594824,"visible":true,"origin":"","legend":"\u003cp\u003eArrangement of the \u003cem\u003esilvana\u003c/em\u003e-beds in the Emerberg-Tautschbuch region redrawn after Schwarz (1913).\u003c/p\u003e","description":"","filename":"Fig.3.jpg","url":"https://assets-eu.researchsquare.com/files/rs-2752635/v1/613b30572d0321146355429a.jpg"},{"id":36275137,"identity":"ca30e24a-6ac6-4d8e-8cd5-6caddc4fed37","added_by":"auto","created_at":"2023-04-25 13:17:34","extension":"jpg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":906884,"visible":true,"origin":"","legend":"\u003cp\u003eDendogram Dice-Index.\u003c/p\u003e","description":"","filename":"Fig.4.jpg","url":"https://assets-eu.researchsquare.com/files/rs-2752635/v1/e9144c575e3ae9fe5fb3aabd.jpg"},{"id":36274229,"identity":"f5632f56-ba4b-4075-8a29-50d98974c31b","added_by":"auto","created_at":"2023-04-25 13:09:34","extension":"jpg","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":805777,"visible":true,"origin":"","legend":"\u003cp\u003eDendogram Kulczynski-Index.\u003c/p\u003e","description":"","filename":"Fig.5.jpg","url":"https://assets-eu.researchsquare.com/files/rs-2752635/v1/072e63ff8998dd9217417d54.jpg"},{"id":36275135,"identity":"2e5cb4db-afca-461c-8d90-bd2f9e8f6517","added_by":"auto","created_at":"2023-04-25 13:17:34","extension":"jpg","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":567947,"visible":true,"origin":"","legend":"\u003cp\u003eDendogram Ochiai-Index\u003c/p\u003e","description":"","filename":"Fig.6.jpg","url":"https://assets-eu.researchsquare.com/files/rs-2752635/v1/2631412e3a45b1132932b09d.jpg"},{"id":36274232,"identity":"b574135b-a734-4154-81ca-ef3e3c9bcada","added_by":"auto","created_at":"2023-04-25 13:09:34","extension":"png","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":4493,"visible":true,"origin":"","legend":"\u003cp\u003eLegend not included with this version\u003c/p\u003e","description":"","filename":"fig.png","url":"https://assets-eu.researchsquare.com/files/rs-2752635/v1/b5cb6f15329838ebce09f64a.png"},{"id":39151978,"identity":"e55cfbb4-37d1-435d-9c3b-9c92b1aa4393","added_by":"auto","created_at":"2023-06-27 08:32:16","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":936451,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-2752635/v1/73bfe631-873f-41bc-bf32-1fd68acb3ab4.pdf"}],"financialInterests":"","formattedTitle":"Paleobiogeography of the land snail communities of the Middle Miocene silvana-beds of Germany","fulltext":[{"header":"1. Introduction","content":"\u003cp\u003eThe \u003cem\u003esilvana\u003c/em\u003e-beds (Middle Miocene) are the main component of the Upper Freshwater Molasse (\u0026ldquo;Obere S\u0026uuml;\u0026szlig;wassermolasse\u0026rdquo; or OSM) unit of the North Alpine Foreland Basin (NAFB) in southern Germany. A large quantity of often well-preserved continental mollusks, especially land snails, have been recovered from these beds, especially during the late 19th and early 20th centuries, being stored in several local and international museum collections. In spite of that, the \u003cem\u003esilvana\u003c/em\u003e-beds have only rarely been subject to systematic excavations or intensive studies. After the pioneering works of Sandberger (1870\u0026ndash;1875), Wenz (1923\u0026ndash;1930) and others, thorough accounts of the \u003cem\u003esilvana\u003c/em\u003e-beds molluscan faunas have only more recently started to appear, although still largely based on historical museum collections (Table\u0026nbsp;1).\u003c/p\u003e \u003cp\u003eOur research group has been steadily collecting information on these fossil mollusks, revising and improving knowledge about them. As such, following the lines of Harzhauser \u0026amp; Mandic (\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2008\u003c/span\u003e) and H\u0026ouml;ltke et al. (\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), our objective here was to gather all information about the land snail communities of the many different outcrops of the \u003cem\u003esilvana\u003c/em\u003e-beds and subject it to a paleobiogeographical analysis. In order to reach this goal, a thorough comparison of these snail faunas was conducted, using similarity indices and follow-up cluster analyses.\u003c/p\u003e"},{"header":"2. The Silvana-beds","content":" \u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003e2.1. Historical background\u003c/h2\u003e \u003cp\u003eDebates regarding the \u003cem\u003esilvana\u003c/em\u003e-beds were exceedingly common in the geological literature of the 19th and early 20th centuries (good examples are Fraas \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e1882\u003c/span\u003e and Rollier \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e1900\u003c/span\u003e). One of the main issues revolved around who first introduced this name and when.\u003c/p\u003e \u003cp\u003eSimply enough, the \u003cem\u003esilvana\u003c/em\u003e-beds are named after the land snail \u003cem\u003ePalaeotachea silvana\u003c/em\u003e (Klein, \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e1853\u003c/span\u003e) (Schad \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e1908\u003c/span\u003e). According to Fraas (\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1888\u003c/span\u003e), the original material of \u003cem\u003eP. silvana\u003c/em\u003e came from M\u0026ouml;rsingen and Mundingen, but the label of the only syntype known names Zwiefalten (near M\u0026ouml;rsingen) as type locality (Salvador et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2016a\u003c/span\u003e). Despite being one of the most frequent fossil land snails in OSM deposits, in the species original description, Klein (\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e1853\u003c/span\u003e) reported that it was fairly rare (\u0026ldquo;seltener\u0026rdquo;, in the original), so he did not mention any possibility of using it as an index fossil.\u003c/p\u003e \u003cp\u003eHowever, the species identity was also up to debate, which caused confusion in the geological literature. Prior to Klein (\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e1853\u003c/span\u003e) and for some time afterwards, the specimens from these beds were identified as \u003cem\u003ePalaeotachea sylvestrina\u003c/em\u003e (Schlotheim, \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e1820\u003c/span\u003e) (e.g., Dunker \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e1848\u003c/span\u003e; Miller \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e1871\u003c/span\u003e), which is now understood to be a separate, younger species (H\u0026ouml;ltke \u0026amp; Rasser \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Fraas (\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e1869\u003c/span\u003e) first proposed \u003cem\u003eP. sylvestrina\u003c/em\u003e as an index fossil for the \u0026ldquo;Hohenmemmiger Kalke\u0026rdquo; (Hohenmemmingen\u0026rsquo;s limestone). Miller (\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e1871\u003c/span\u003e) then used the term \u003cem\u003esylvestrina\u003c/em\u003e-beds for the OSM sediments in Hochstr\u0026auml;\u0026szlig;, near Ulm, without mention to \u003cem\u003eP. silvana\u003c/em\u003e. Sandberger (1870\u0026ndash;1875) then listed \u003cem\u003eP. silvana\u003c/em\u003e from a several localities in the NAFB, but without considering its use as an index fossil. Quenstedt (1876), however, continued to use only the name \u003cem\u003eP. sylvestrina\u003c/em\u003e as well as the term \u0026ldquo;Sylvestrinenkalke\u0026rdquo; (\u003cem\u003esylvestrina\u003c/em\u003e-limestone). Deffner \u0026amp; Fraas (\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e1877\u003c/span\u003e) then listed both species (\u003cem\u003eP. silvana\u003c/em\u003e and \u003cem\u003eP. sylvestrina\u003c/em\u003e) in their Tertiary chapter. Finally, Probst (\u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e1879\u003c/span\u003e: 265\u0026ndash;266) was the first to indicate \u003cem\u003eP. silvana\u003c/em\u003e as an index fossil for the OSM; the identification of his material was done by Sandberger. Quenstedt (\u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e1884\u003c/span\u003e) still retained the name \u003cem\u003esylvestrina\u003c/em\u003e-beds, arguing that it was the common usage among geologists and that this species was known as \u003cem\u003eP. silvana\u003c/em\u003e in the Swabian region. Fraas (\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e1882\u003c/span\u003e) simply considered \u003cem\u003eP. silvana\u003c/em\u003e as synonym of \u003cem\u003eP. sylvestrina\u003c/em\u003e. However, Engel (\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e1883\u003c/span\u003e: 274) named the unit \u003cem\u003esilvana\u003c/em\u003e-beds due to the index fossil \u003cem\u003eP. silvana\u003c/em\u003e; in his words, a very practical name (\u0026ldquo;sehr praktischer Namen\u0026rdquo;).\u003c/p\u003e \u003cp\u003eZittel (\u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e1885\u003c/span\u003e) then listed both species, \u003cem\u003eP. silvana\u003c/em\u003e and \u003cem\u003eP. sylvestrina\u003c/em\u003e, for the OSM, but without using the terms \u003cem\u003esilvana\u003c/em\u003e- or \u003cem\u003esylvestrina\u003c/em\u003e-beds. Fraas (\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1888\u003c/span\u003e) finally adopted the term \u003cem\u003esilvana\u003c/em\u003e-limestone (\u0026ldquo;Silvanakalk\u0026rdquo;), but still considered \u003cem\u003eP. silvana\u003c/em\u003e and \u003cem\u003eP. sylvestrina\u003c/em\u003e to represent the same species.\u003c/p\u003e \u003cp\u003eIn summary, from the first use of \u003cem\u003eP. silvana\u003c/em\u003e as index fossil (Probst, \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e1879\u003c/span\u003e), through the naming of the unit as \u003cem\u003esilvana\u003c/em\u003e-beds (Engel, \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e1883\u003c/span\u003e) and onwards, several different opinions remained in the literature. These concerned both the unit\u0026rsquo;s name and the validity of P. \u003cem\u003esilvana\u003c/em\u003e as a separate species. Nevertheless, by the beginning of the 20th century, the name \u003cem\u003esilvana\u003c/em\u003e-beds prevailed (e.g., Rollier \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e1900\u003c/span\u003e, \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e1903\u003c/span\u003e; Miller \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e1901\u003c/span\u003e, \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e1903\u003c/span\u003e; Jooss \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e1915\u003c/span\u003e; Gottschick \u0026amp; Wenz \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e1916\u003c/span\u003e). Still, while some authors started to focus on describing lithological subunits within the \u003cem\u003esilvana\u003c/em\u003e-beds (Schad \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e1908\u003c/span\u003e for the Ehingen/Donau region; Schwarz \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e1913\u003c/span\u003e, for the Emerberg/Tautschbuch region), another issues arose: the stratigraphical position and age of the beds. Researchers considered two main possibilities, either Upper Oligocene (Rollier \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e1900\u003c/span\u003e) or Upper Miocene (Miller \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e1901\u003c/span\u003e; Jooss \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e1915\u003c/span\u003e; Dietrich \u0026amp; Kautsky, \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e1920\u003c/span\u003e). Finally, Wenz (\u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e1920\u003c/span\u003e) summarized the then current knowledge and published a classification of the \u003cem\u003esilvana\u003c/em\u003e beds, dividing them in three sections based on the land snail fauna (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e3\u003c/span\u003e; see also H\u0026ouml;ltke \u0026amp; Rasser \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2016\u003c/span\u003e) and placing it in the Upper Miocene (Tortonian). Furthermore, Wenz (\u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e1921\u003c/span\u003e) separated the units into the older \u003cem\u003esilvana\u003c/em\u003e-beds and the younger \u003cem\u003esylvestrina\u003c/em\u003e-beds. The lowest section of the \u003cem\u003esilvana\u003c/em\u003e-beds received a special name, \u0026ldquo;Flammenmergel\u0026rdquo; or \u0026ldquo;flame marl\u0026rdquo; (Wenz \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e1924\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eIn the decades that followed, publications about the \u003cem\u003esilvana\u003c/em\u003e-beds became much scarcer, with some accounts of fossil snails (Gottschick \u0026amp; Wenz \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e1927\u003c/span\u003e; Schlickum \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e1976\u003c/span\u003e) and regional geological works (Haag \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e1960\u003c/span\u003e; Z\u0026ouml;belein \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e1977\u003c/span\u003e). Only much later, the \u003cem\u003esilvana\u003c/em\u003e-beds started to draw renewed attention (Schweigert \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e1996\u003c/span\u003e; Esu \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Doppler et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). Esu (\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e1999\u003c/span\u003e) was responsible for placing the \u003cem\u003esilvana\u003c/em\u003e-beds within the Neogene Mammal zone MN5 of the Middle Miocene); hence, these beds predate the N\u0026ouml;rdlinger Ries and Steinheim am Albuch meteorite events. Starting with Sach (\u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e1999\u003c/span\u003e), the molluscan fauna of the \u003cem\u003esilvana\u003c/em\u003e-beds began to be re-examined (see publications listed in Table\u0026nbsp;1). Finally, H\u0026ouml;ltke \u0026amp; Rasser (\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2016\u003c/span\u003e) revised the \u003cem\u003ePalaeotachea\u003c/em\u003e species from the German Miocene and discussed the implications for stratigraphy.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003e2.2. Stratigraphy and lithology\u003c/h2\u003e \u003cp\u003eAs explained above, the \u003cem\u003esilvana\u003c/em\u003e-beds are a unit within the OSM in the North Alpine Foreland Basin (for details about this basin, see K\u0026auml;lin \u0026amp; Kempf \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2009\u003c/span\u003e), defined by the presence of the index fossil \u003cem\u003ePalaeotachea silvana\u003c/em\u003e, a terrestrial pulmonate gastropod. The strata underlying the \u003cem\u003esilvana\u003c/em\u003e-beds are region-dependant and can be: the \u0026ldquo;Helicidenmergel\u0026rdquo; (\u0026ldquo;helicid-marl\u0026rdquo;), the Upper Brackish Water Molasse (\u0026ldquo;Brackwassermolasse\u0026rdquo;) or the Upper Jusassic Malm. Immediately above the \u003cem\u003esilvana\u003c/em\u003e-beds are the \u003cem\u003esilvestrina\u003c/em\u003e-beds (H\u0026ouml;ltke \u0026amp; Rasser \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2016\u003c/span\u003e) or, where the latter is absent, Pleistocene gravel. The \u003cem\u003esilvana\u003c/em\u003e-beds are freshwater sediments, consisting of limestone, marl, silt and also partly with a sandy component. Naturally, they also include several fossils, especially gastropods. In particular, the deposits of the Emerberg/Tauschbuch region at the southern margin of the Swabian Alb (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e1\u003c/span\u003e) are a hotspot of diversity and preservation of land snails (H\u0026ouml;ltke et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eWenz (\u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e1920\u003c/span\u003e, \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e1921\u003c/span\u003e, \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e1924\u003c/span\u003e) divided the \u003cem\u003esilvana\u003c/em\u003e-beds in three different subunits based mainly on their land snail fauna (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Despite some criticisms (Jooss \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e1923\u003c/span\u003e; Z\u0026ouml;belein \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e1977\u003c/span\u003e) regarding wider stratigraphic distribution of some species, the structuring of the \u003cem\u003esilvana\u003c/em\u003e-beds \u003cem\u003esensu\u003c/em\u003e Wenz is a practical tool, especially when dealing with the usually incomplete information available for most localities. Furthermore, more region-specific descriptions and subdivisions of the \u003cem\u003esilvana\u003c/em\u003e-beds were published by Schad (\u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e1908\u003c/span\u003e), Schwarz (\u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e1913\u003c/span\u003e) (see Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e3\u003c/span\u003e), Haag (\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e1960\u003c/span\u003e) and Z\u0026ouml;belein (\u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e1977\u003c/span\u003e); the latter also provided a critical overview about the works of the former three authors.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e"},{"header":"3. Material And Methods","content":"\u003cp\u003eWe compiled literature data on all localities belonging to the \u003cem\u003esilvana\u003c/em\u003e-beds, using the most up to date works available. Moreover, the coeval localities from the neighboring Bad Urach volcanic region of the Swabian Alb (which lies just outside the NAFB) were included. As a compromise between the scarcity of data for several localities and a meaningful number for the analyses using similarity indices, we only considered those localities with at least six species of land snails. The chosen localities are listed in Table\u0026nbsp;1.\u003c/p\u003e \u003cp\u003eIn some cases, the literature available dates from the early 20th century, so we corrected the species lists they presented regarding minor taxonomic issues (synonymies and new names) according to the more recent literature (Table\u0026nbsp;1). In a few cases, literature data could be complemented with material from museum collections, namely: Staatliches Museum f\u0026uuml;r Naturkunde Stuttgart (SMNS; Stuttgart, Germany), Bayerische Staatssammlung f\u0026uuml;r Pal\u0026auml;ontologie und Geologie (BSPG; Munich, Germany), and Department of Geosciences of the Eberhard Karls Universit\u0026auml;t T\u0026uuml;bingen (GPIT; T\u0026uuml;bingen, Germany; formerly Geologisch-Pal\u0026auml;ontologisches Institut T\u0026uuml;bingen).\u003c/p\u003e \u003cp\u003eThe resulting list of land snail species for each locality was then subjected to a cluster analysis using four distinct similarity indices with \u0026ldquo;Paired group (UPGMA)\u0026rdquo; algorithm. The indices used for the presence-absence data of the species were: S\u0026oslash;rensen-Dice (Dice \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1945\u003c/span\u003e; S\u0026oslash;rensen \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e1948\u003c/span\u003e), Kulczynski (\u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e1927\u003c/span\u003e), Ochiai (\u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e1957\u003c/span\u003e), and Simpson (\u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e1943\u003c/span\u003e, \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e1960\u003c/span\u003e). This was done to test for stability and consistency of the groupings in the resulting dendrograms. In this step, we removed those species endemic to a single site. The analyses were conducted with 10,000 bootstrap replicates to test for the robustness of the groups (robustness estimates, in percentages, are shown at each node of the dendograms). The analyses were carried out in PAST v.3.16 (Hammer et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). Finally, we searched for possible common land snail micro-communities, that is, a constant assemblage of some species that could be found on more than one locality. However, the cluster analysis results were inconsistent, possibly due to the small sample size of most localities or slight differences in stratigraphical position between geographically close localities. Therefore, this step is omitted in the present article.\u003c/p\u003e"},{"header":"4. Results","content":"\u003cp\u003eA total of 28 localities were used in the present work (Table\u0026nbsp;1), containing together 51 different land snail species. The localities with the highest number of species are M\u0026ouml;rsingen and Zwiefaltendorf, with 36 each.\u003c/p\u003e \u003cp\u003eThe dendrograms resulting from the cluster analyses (Figs.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e\u0026ndash;7) share similarities and differences. To conduct comparisons between them, only clusters with a similarity index of ca. 50% were taken into account (for the Simpson index, subgroups with ca. 65% similarity were explored; Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e6\u003c/span\u003e). The clusters were numbered in Figs.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e3\u003c/span\u003e\u0026ndash;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e6\u003c/span\u003e and these numbers are used throughout the text. The robustness estimates from the bootstrap are usually low for the large clusters, but can achieve high values for some locality pairs (e.g., M\u0026ouml;rsingen and Zwiefaltendorf). Low values normally indicate that the cluster is not well supported; however, in the present case the low values might be exacerbated due to the small sample size of several terminals (localities) and the thus disproportionate importance of the few species-rich terminals inside the clusters. Robustness estimates values are especially low for the Simpson index, but the results of this index are also carefully discussed herein as it is deemed very appropriate for paleontological samples in which there are large differences in species richness between localities (Simpson \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e1943\u003c/span\u003e, \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e1960\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eThe Bavarian localities Riedensheim, Sandelzhausen and Adelschlag-Fasanerie form one reasonably well-supported cluster in all indices except for Simpson, which includes other localities (Figs.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e\u0026ndash;7: clusters D1, K1, O1, S1). In all dendrograms, there is one large cluster containing most of the Baden-W\u0026uuml;rttemberg deposits located at the Southern and Eastern margin of the Swabian Alb (Figs.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e\u0026ndash;7: clusters D2, K2, O3, S2). In all cases, the Bavarian locality G\u0026uuml;ndlkofen is clustered with the Austrian Pf\u0026auml;nder region (Figs.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e\u0026ndash;7: clusters D6, K5, O6, S4). The localities Georgensgm\u0026uuml;nd and Mundingen form one cluster by the S\u0026oslash;rensen-Dice and Kulczynski indices, but include Ehingen/Donau by the Ochiai index (Figs.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e\u0026ndash;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e6\u003c/span\u003e: clusters D5, K4, O4); this cluster is absent in the Simpson index dendrogram. In all dendrograms, the localities D\u0026auml;chingen, Randecker Maar, Hengen and Oggenhausen are clustered together (Figs.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e\u0026ndash;7: clusters D3, K3, O2, S3), but by the Ochiai index, it also contains the localities Daugendorf and Andelfingen. There is an additional large cluster identified by the S\u0026oslash;rensen-Dice index (D4), containing four localities that are scattered through other clusters according to the other indices. The locality Baach is separated from all others in all four indices and the position of the locality Vermes is extremely variable.\u003c/p\u003e \u003cp\u003eIn the dendrogram resulting from Simpson index (Fig.\u0026nbsp;7), there are five pairs of localities with 100% similarity. This is because this index is especially good in detecting nestedness of samples, that is, if one community contains the same species as another community and a few extra ones, the latter is nested within the former (index value equals one).\u003c/p\u003e"},{"header":"5. Discussion","content":"\u003cp\u003eThe Dice, Kuclczynski and the Ochiai index plots are overall very similar to one another, whereas the Simpson index plot is the most different. Nevertheless, despite these differences, the basic arrangement is still similar enough for meaningful comparison. The similarities of clusters are discussed in further detail below and possible reasons for the observed differences are offered.\u003c/p\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003e5.1. Bavarian cluster\u003c/h2\u003e \u003cp\u003eThe cluster containing the Bavarian localities Riedensheim, Sandelzhausen and Adelschlag-Fasanerie (Figs.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e\u0026ndash;7: clusters D1, K1, O1, S1) can be explained by their geographic position: the three deposits are geographically very close. These localities have also been deemed very similar in previous works (H\u0026ouml;ltke et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Salvador et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2016b\u003c/span\u003e), which have also included Undorf, another closely located deposit. Undorf, however, here is included in the Baden-W\u0026uuml;rttemberg cluster (see below) for all indices except Simpson (Fig.\u0026nbsp;7). By the latter, Undorf is featured together with the three localities above and also the Swiss deposits of Vermes.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec9\" class=\"Section2\"\u003e \u003ch2\u003e5.2. Vermes\u003c/h2\u003e \u003cp\u003eThe only Swiss locality, Vermes, has a very variable position in the dendrograms. The locality is generally well separated from the others, with low similarity values. According to the S\u0026oslash;rensen-Dice and Ochiai indices (Figs.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e, \u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e6\u003c/span\u003e), it is more closely related to Baden-W\u0026uuml;rttemberg localities, while the Kulczynski and Simpson indices place it closer to the Bavarian localities. In fact, by the latter index, it is placed well inside the Bavarian group (see above), together with the locality Undorf. The reason for this clustering, despite their geographic distance, could be the high number of species it shares with Undorf (seven: \u003cem\u003eArchaeozonites costatus\u003c/em\u003e, \u003cem\u003eGastrocopta nouletiana\u003c/em\u003e, \u003cem\u003eLeucochroopsis kleini\u003c/em\u003e, \u003cem\u003ePalaeotachea silvana\u003c/em\u003e, \u003cem\u003eTestacella zelli\u003c/em\u003e, \u003cem\u003eTriptychia kleini\u003c/em\u003e, and \u003cem\u003eVitrina suevica\u003c/em\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec10\" class=\"Section2\"\u003e \u003ch2\u003e5.3. Baden-W\u0026uuml;rttemberg cluster\u003c/h2\u003e \u003cp\u003eThis large cluster can be found, with a few differences, in all indices (Figs.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e\u0026ndash;7: clusters D2, K2, O3, S2). Most of the localities are located on the southern margin of the Swabian Alb (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e1\u003c/span\u003e), which is a likely reason behind their similarity in species composition. Dischingen and Hohenmemmingen, in particular, are slightly removed towards the eastern margin of the Alb, but still fairly close.\u003c/p\u003e \u003cp\u003eAn unusual locality within this cluster is Undorf, located in Bavaria. Undorf is often deemed more related to the Bavarian cluster (H\u0026ouml;ltke et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Salvador et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2016b\u003c/span\u003e), which is more reasonable from a paleogeographical point of view. In fact, the results of the Simpson index places Undorf within the Bavaria cluster (Fig.\u0026nbsp;7). The reason for the grouping of Undorf with the Baden-W\u0026uuml;rttemberg localities might be due to the 21 species it shares with the joint localities from the southern margin of the Swabian Alb, because the other indices used herein give more weight to shared species between localities.\u003c/p\u003e \u003cp\u003eOther inconsistencies are the presence of the Bavarian locality Georgensgm\u0026uuml;nd by Simpson index (Fig.\u0026nbsp;7) and the absence of Friedingen (near to Zwiefalten and thus withing the geographical region) according to S\u0026oslash;rensen-Dice index.\u003c/p\u003e \u003cp\u003eThe localities within this cluster may represent the lower and middle \u003cem\u003esilvana\u003c/em\u003e-beds (\u003cem\u003esensu\u003c/em\u003e Wenz \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e1920\u003c/span\u003e, \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e1921\u003c/span\u003e, \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e1924\u003c/span\u003e) (see also Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e3\u003c/span\u003e) due to the abundance of the species \u003cem\u003eP. silvana\u003c/em\u003e and \u003cem\u003ePseudochloritis incrassata\u003c/em\u003e, as well as the occurrence of \u003cem\u003ePalaeotachea dentula\u003c/em\u003e in some deposits.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003e5.4. Maar cluster\u003c/h2\u003e \u003cp\u003eThis cluster is made up of \u0026ldquo;geographically mixed\u0026rdquo; localities (Figs.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e\u0026ndash;7: clusters D3, K3, O2, S3), with most localities \u0026ldquo;scattered\u0026rdquo; through Baden-W\u0026uuml;rttemberg state. According to S\u0026oslash;rensen-Dice and Simpson indices, this clusters consist of Randecker Maar, Hengen, D\u0026auml;chingen and Oggenhausen (Figs.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e, 7: clusters D3, S3), which share overall five species (\u003cem\u003eArchaeozonites costatus\u003c/em\u003e, \u003cem\u003ePalaeotachea renevieri\u003c/em\u003e, \u003cem\u003eP. silvana\u003c/em\u003e, \u003cem\u003ePomatias conica\u003c/em\u003e, and \u003cem\u003ePseudochloritis incrassata\u003c/em\u003e). The Kulzynski index also includes Andelfingen (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e5\u003c/span\u003e: cluster K3), resulting in overall three species in common (\u003cem\u003eA. costatus, P. silvana, P. incrassata\u003c/em\u003e). Finally, the Ochiai index includes both Andelfingen and Daugendorf (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e6\u003c/span\u003e: cluster O2), resulting in only two species in common \u003cem\u003e(P. silvana, P. incrassata\u003c/em\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eThe pairing of the localities Randeck Maar and Hengen in all indices except Simpson, could be related to the fact that both had a maar lake paleoenvironment. D\u0026auml;chingen and Oggenhausen had very different paleoenvironments, but the few species shared with the others may have influenced this clustering. The reasons for the inclusion of Daugendorf and Andelfingen are unclear. They have only the two most typical species in common with the other four localities (\u003cem\u003eP. silvana\u003c/em\u003e \u0026amp; \u003cem\u003eP. incrassata\u003c/em\u003e). According to the presence of \u003cem\u003eP. incrassata\u003c/em\u003e the reason for this clustering maybe the same stratigraphic position of deposits named above (Middle \u003cem\u003esilvana\u003c/em\u003e-beds sensu Wenz \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e1924\u003c/span\u003e) (see also Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec12\" class=\"Section2\"\u003e \u003ch2\u003e5.5. Cluster D4\u003c/h2\u003e \u003cp\u003eThe cluster D4 appears exclusively on the dendrogram from the S\u0026oslash;rensen-Dice index (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). It only contains deposits located on the southern margin of the Swabian Alb, but not included in the Baden-W\u0026uuml;rtemberg cluster (see above) by this index. In all other indices, the localities forming the D4 cluster are usually part of the Baden-W\u0026uuml;rtemberg cluster. All the localities have only three species in common: \u003cem\u003eLeucochroopsis kleini\u003c/em\u003e, \u003cem\u003ePalaeotachea silvana\u003c/em\u003e, \u003cem\u003ePseudochloritis incrassata\u003c/em\u003e. The reasons for the S\u0026oslash;rensen-Dice index separating this second Swabian Alb cluster are unclear, but may be related to the absence of some key species or to different stratigraphic positions.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003e5.6. Georgensgm\u0026uuml;nd/Mundingen clusters\u003c/h2\u003e \u003cp\u003eIn all indices except for Simpson, Mundingen (part of the Ehingen/Donau district) in Baden-W\u0026uuml;rttemberg forms one cluster with the Bavarian locality Georgensgm\u0026uuml;nd (Figs.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e\u0026ndash;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e6\u003c/span\u003e: clusters D5, K4 and O4). Despite their different geographic positions, the two localities have four species in common (\u003cem\u003eApula coarctata, Janulus supracostatus\u003c/em\u003e, \u003cem\u003eP. silvana\u003c/em\u003e, and \u003cem\u003ePseudochloritis incrassata\u003c/em\u003e). The Ochiai index also includes the locality Ehingen/Donau, which is close to Mundingen. Nevertheless, the three deposits only share the two typical \u003cem\u003esilvana\u003c/em\u003e-beds species (\u003cem\u003eP. silvana\u003c/em\u003e and \u003cem\u003ePseudochloritis incrassata\u003c/em\u003e). The reason for this clustering is also unclear, but could be related to similar stratigraphic positions within the \u003cem\u003esilvana\u003c/em\u003e-beds or to the influence of paleoenvironmental factors.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec14\" class=\"Section2\"\u003e \u003ch2\u003e5.7. G\u0026uuml;ndlkofen/Pf\u0026auml;nder cluster\u003c/h2\u003e \u003cp\u003eThe Bavarian locality G\u0026uuml;ndlkofen and the Austrian deposit Pf\u0026auml;nder (near Bregenz) form one cluster, quite isolated from all others (Figs.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e3\u003c/span\u003e\u0026ndash;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e6\u003c/span\u003e: D6, K5, O5, S4). The two localities have four species in common: \u003cem\u003eP. silvana, Pseudidyla moersingensis, Pseudochloritis incrassata, Triptychia teutonica\u003c/em\u003e. The reasons for the clustering might be either geographic or stratigraphic, as G\u0026uuml;ndlkofen is deemed to be younger than the other Bavarian localities.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec15\" class=\"Section2\"\u003e \u003ch2\u003e5.8. Baach\u003c/h2\u003e \u003cp\u003eThe locality Baach (part of the Zwiefalten district) is located on the southern margin of the Swabian Alb in Baden-W\u0026uuml;rttemberg, but it is completely isolated from all other localities in all four cases. All the species occurring in Baach can also found in the deposits from the Swabian Alb. The most important features of Baach are the missing of of \u003cem\u003ePseudochloritis incrassata\u003c/em\u003e and the presence of \u003cem\u003ePalaeotachea dentula\u003c/em\u003e. So the reason for the separation may be due to the stratigraphic position representing the lower \u003cem\u003esilvana\u003c/em\u003e-beds sensu Wenz \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e1924\u003c/span\u003e (see also Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e"},{"header":"6. Conclusion","content":"\u003cp\u003eThe different clusters in the present biogeographic analyses can be largely explained by their geographic position, like the Bavarian and Baden-W\u0026uuml;rttemberg clusters. Some inconsistencies in the composition of the clusters might be explained by different stratigraphic positions within the \u003cem\u003esilvana\u003c/em\u003e-beds (Georgensgm\u0026uuml;nd/Mundingen and G\u0026uuml;ndlkofen/Pf\u0026auml;nder clusters), distinct paleoenvironmental conditions influencing the faunal composition (Maar cluster) or absence of key species (maybe Baach). However, with the scarce data presently available, it is difficult to consider the likelihood of such hypotheses.\u003c/p\u003e \u003cp\u003eAt present, the \u003cem\u003esilvana\u003c/em\u003e-beds (and also the \u003cem\u003esylvestrina\u003c/em\u003e-beds) have not been defined according to international stratigraphical guidelines, thus remaining an informal lithostratigraphic unit. The only distinct feature of the \u003cem\u003esilvana\u003c/em\u003e-beds is the presence of the land snail \u003cem\u003eP. silvana\u003c/em\u003e, which is also true for the \u003cem\u003esylvestrina\u003c/em\u003e-beds and \u003cem\u003eP. sylvestrina\u003c/em\u003e. Perhaps it would be sensible to define it as biostratigraphic zone, in the lines of the Mesozoic ammonoid zones. In this way, the coeval sediments from the nearby Bad Urach volcano region (e.g., Hengen), which is just outside the NAFB and bear the same land snail fauna, could be included without problems in the \u003cem\u003esilvana\u003c/em\u003e-beds. However, for detailed stratigraphic research on the \u003cem\u003esilvana\u003c/em\u003e-beds, good outcrops/profiles would be necessary, which are unfortunately unavailable in the present.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003eFunding: None.\u003c/p\u003e\n\u003cp\u003eConflicts of interest/Competing interests: There are no conflicts of interest.\u003c/p\u003e\n\u003cp\u003eAvailability of data and material: The material is stored in The Staatlichen Museum f\u0026uuml;r Naturkunde Stuttgart.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eCode availability: Not applicable.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eAuthors\u0026apos; contributions: Not applicable.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eClessin, S. (19111). Die Conchylien der obermiocaenen Ablagerung von Undorf. \u003cem\u003eBerichte des naturwissenchaftlichen Vereins zu Regensburg\u003c/em\u003e, 13: 1\u0026ndash;13.\u003c/li\u003e\n\u003cli\u003eDeffner, C. \u0026amp; Fraas, O. (1877). \u003cem\u003eBegleitworte zur geognostischen Specialkarte von W\u0026uuml;rttemberg. \u003c/em\u003e\u003cem\u003eAtlasbl\u0026auml;tter Bopfingen und Ellenberg\u003c/em\u003e. Kohlhammer.\u003c/li\u003e\n\u003cli\u003eDice, L.R. (1945). Measures of the Amount of Ecologic Association Between Species. \u003cem\u003eEcology\u003c/em\u003e. 26 (3): 297\u0026ndash;302. http://dx.doi.org/10.2307/1932409\u003c/li\u003e\n\u003cli\u003eDietrich, W.O. \u0026amp; Kautsky, F. (1920). Die Altersbeziehungen der schw\u0026auml;bischen und schweizerischen oberen Meeresmolasse und des Terti\u0026auml;rs am S\u0026uuml;drand der Schw\u0026auml;bischen Alb. \u003cem\u003eCentralblatt f\u0026uuml;r Mineralogie, Geologie und Pal\u0026auml;ontologie\u003c/em\u003e, 1920: 243\u0026ndash;253.\u003c/li\u003e\n\u003cli\u003eDoppler, G., Heissig, K. \u0026amp; Reichenbacher, B. (2005). Die Gliederung des Terti\u0026auml;rs im s\u0026uuml;ddeutschen Molassebecken. Newsletter Stratigraphy, 41(1-3): 359\u0026ndash;375. DOI: 10.1127/0078-0421/2005/0041-0359\u003c/li\u003e\n\u003cli\u003eDunker, W. (1848). Ueber die in der Molasse bei G\u0026uuml;nzburg unfern Ulm vorkommenden Conchylien und Pflanzenreste. \u003cem\u003ePalaeontographica\u003c/em\u003e, 1:155\u0026ndash;168.\u003c/li\u003e\n\u003cli\u003eEhrat, H. \u0026amp; Jooss, C.H. (1921). Das Alter der vulkanischen Tuffe im Kirchheim-Uracher Gebiet und im Hegau. \u003cem\u003eGeologische und Pal\u0026auml;ontologische Mitteilungen\u003c/em\u003e, 1: 1\u0026ndash;8.\u003c/li\u003e\n\u003cli\u003eEngel, T. (1883). \u003cem\u003eGeognostischer Wegweiser durch W\u0026uuml;rttemberg\u003c/em\u003e. 326 pp. Stuttgart (Schweizerbart).\u003c/li\u003e\n\u003cli\u003eEsu, D. (1999). \u003cem\u003eContribution to the knowledge of Neogene climatic changes in western and central Europe by means of non-marine molluscs\u003c/em\u003e. Agusti, J., Rook, L. \u0026amp; Andrews, P. (Eds.), \u003cem\u003eHominid Evolution and Climatic Change in Europe, Vol. 1. The Evolution of Neogene Terrestrial Ecosystems in Europe\u003c/em\u003e, (pp. 328\u0026ndash;354). Cambridge University Press.\u003c/li\u003e\n\u003cli\u003eFraas, O. (1869). \u003cem\u003eBegleitworte zur geognostischen Specialkarte von W\u0026uuml;rttemberg. 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Neue Helicellinen aus schw\u0026auml;bischen Silvanaschichten. \u003cem\u003eArchiv f\u0026uuml;r Molluskenkunde\u003c/em\u003e, 59: 147\u0026ndash;149.\u003c/li\u003e\n\u003cli\u003eHaag, H.W. (1960). Die Geologie des Blattes Zweifalten (Nr. 7722) 1 : 25 000 (Stratigraphie und Tektonik der Zwiefalter Alb). \u003cem\u003eArbeiten aus dem Geologisc Pal\u0026auml;ontologischen Institut der Technischen Hochschule Stuttgart, N.F\u003c/em\u003e., 28: 1\u0026ndash;121.\u003c/li\u003e\n\u003cli\u003eHammer, \u0026Oslash;., Harper, D.A.T. \u0026amp; Ryan, P.D. (2001). PAST: Paleontological statistics software package for education and data analysis. \u003cem\u003ePalaeontologia Electronica\u003c/em\u003e, 4(1): 1\u0026ndash;9.\u003c/li\u003e\n\u003cli\u003eHarzhauser, M. \u0026amp; Mandic, O. (2008). Neogene lake systems of Central and South-Eastern Europe: Faunal diversity, gradients and interrelations. \u003cem\u003ePalaeogeography, Palaeoclimatology, Palaeoecology\u003c/em\u003e, 260: 417\u0026ndash;434. http://dx.doi.org/10.1016/j.palaeo.2007.12.013\u003c/li\u003e\n\u003cli\u003eH\u0026ouml;ltke, O. \u0026amp; Rasser M.W. (2016). The \u003cem\u003ePalaeotachea\u003c/em\u003e complex (Gastropoda: Pulmonata) in the Miocene of SW Germany: a morphometric approach. \u003cem\u003eJournal of Conchology\u003c/em\u003e, 42(4): 239\u0026ndash;256.\u003c/li\u003e\n\u003cli\u003eH\u0026ouml;ltke, O., Salvador, R.B. \u0026amp; Rasser, M.W. (2016). Paleobiogeography of Early/Middle Miocene terrestrial gastropods in Central Europe: an approach using similarity indices. \u003cem\u003ePalaeogeography, Palaeoclimatology, Palaeoecology\u003c/em\u003e, 461: 224\u0026ndash;236. http://dx.doi.org/10.1016/j.palaeo.2016.08.027\u003c/li\u003e\n\u003cli\u003eH\u0026ouml;ltke, O., Salvador, R.B. \u0026amp; Rasser, M.W. (2018). Miocene continental gastropods from the southern margin of the Swabian Alb (Baden-W\u0026uuml;rttemberg, SW Germany). \u003cem\u003eNeues Jahrbuch f\u0026uuml;r Geologie und Pal\u0026auml;ontologie, Abhandlungen,\u003c/em\u003e 287(1): 17\u0026ndash;44. http://dx.doi.org/10.1127/njgpa/2018/0704\u003c/li\u003e\n\u003cli\u003eJooss, C.H. (1910). Binnenconchylienfauna aus dem Obermioz\u0026auml;n des Pf\u0026auml;nders bei Bregenz am Bodensee. \u003cem\u003eNachrichtsblatt der Deutschen Malakozoologischen Gesellschaft\u003c/em\u003e, \u003cstrong\u003e42\u003c/strong\u003e: 19\u0026ndash; 29.\u003c/li\u003e\n\u003cli\u003eJooss, C.H. (1915). Zur Altersfrage der S\u0026uuml;\u0026szlig;wasserablagerungen bei der Ruggburg am\u003c/li\u003e\n\u003cli\u003ePf\u0026auml;nder bei Bregenz. \u003cem\u003eCentralblatt f\u0026uuml;r Mineralogie, Geologie und Palaeontologie\u003c/em\u003e, 1915: 62\u0026ndash;64.\u003c/li\u003e\n\u003cli\u003eJooss, C.H. (1923). Die Schneckenfauna der s\u0026uuml;ddeutsch-schweizerischen Helicidenmergel und ihre Bedeutung f\u0026uuml;r die Altersbestimmung der letzteren. \u003cem\u003eNeues Jahrbuch f\u0026uuml;r Geologie und Pal\u0026auml;ontologie\u003c/em\u003e, 49: 185 ̶ 211.\u003c/li\u003e\n\u003cli\u003eK\u0026auml;lin, D., \u0026amp; Kempf, O. (2009). High-resolution stratigraphy from the continental record of the Middle Miocene Northern Alpine Foreland Basin of Switzerland. \u003cem\u003eNeues Jahrbuch f\u0026uuml;r Geologie und Pal\u0026auml;ontologie, Abhandlungen\u003c/em\u003e, 254: 177\u0026ndash;235. http://dx.doi.org/10.1127/0077-7749/2009/0010\u003c/li\u003e\n\u003cli\u003eKlein, A. von (1846). Conchylien der S\u0026uuml;sswasserkalkformation W\u0026uuml;rttembergs. J\u003cem\u003eahreshefte des Vereins f\u0026uuml;r vaterl\u0026auml;ndische Naturkunde in W\u0026uuml;rttemberg\u003c/em\u003e, 2: 60\u0026ndash;116.\u003c/li\u003e\n\u003cli\u003eKlein, A. von (1853). Conchylien der S\u0026uuml;\u0026szlig;wasserkalkformation W\u0026uuml;rttembergs. \u003cem\u003eJahreshefte des Vereins f\u0026uuml;r vaterl\u0026auml;ndische Naturkunde in W\u0026uuml;rttemberg\u003c/em\u003e, 9: 203\u0026ndash;223.\u003c/li\u003e\n\u003cli\u003eKulczynski, S. (1927). Die Pflanzenassoziationen der Pieninen. \u003cem\u003eBulletin International de l\u0026rsquo;Academie Polonaise des Sciences et des Lettres, Classe des Sciences Mathematiques et Naturelles B\u003c/em\u003e, 1927: 57\u0026ndash;203.\u003c/li\u003e\n\u003cli\u003eMaassen, W.J.M. (2003). Notes on terrestrial molluscs of the island of Sulawesi. 3. The genera Palaina, Arinia and Opisthostoma (Gastropoda, Prosobranchia, Diplommatinidae), with descriptions of a dozen new taxa. \u003cem\u003eBasteria\u003c/em\u003e, 67: 47\u0026ndash; 63.\u003c/li\u003e\n\u003cli\u003eMiller, K. (1871). Das Terti\u0026auml;r am Hochstr\u0026auml;ss. \u003cem\u003eJahreshefte des Vereins f\u0026uuml;r vaterl\u0026auml;ndische Naturkunde in W\u0026uuml;rttemberg,\u003c/em\u003e 27: 272\u0026ndash;292.\u003c/li\u003e\n\u003cli\u003eMiller, K. (1901). Zum Alter des Sylvanakalkes. \u003cem\u003eCentralblatt f\u0026uuml;r Mineralogie, Geologie und Palaeontologie\u003c/em\u003e, 1901: 129\u0026ndash;133.\u003c/li\u003e\n\u003cli\u003eMiller, K. (1903). Zu Rollier, das Alter des Sylvanakalks.-\u003cem\u003eCentralblatt f\u0026uuml;r Mineralogie, Geologie und Palaeontologie\u003c/em\u003e, 1903: 141\u0026ndash;144.\u003c/li\u003e\n\u003cli\u003eNekola, J.C. \u0026amp; Coles, B.F. (2010). Pupillid Land Snails of Eastern North America. \u003cem\u003eAmerican Malacological Bulletin\u003c/em\u003e, 28(2): 29\u0026ndash;57. http://dx.doi.org/10.4003/006.028.0221\u003c/li\u003e\n\u003cli\u003eOchiai, A. (1957). Zoographic studies on the soleoid fishes found in Japan and its neighboring regions. \u003cem\u003eBulletin of the Japanese Society of Scientific Fisheries\u003c/em\u003e, 22: 526\u0026ndash;530.\u003c/li\u003e\n\u003cli\u003eProbst, J. (1879). Verzeichnis der Fauna und Flora der Molasse im w\u0026uuml;rttembergischen\u003c/li\u003e\n\u003cli\u003eOberschwaben. \u003cem\u003eJahreshefte des Vereins f\u0026uuml;r vaterl\u0026auml;ndische Naturkunde in W\u0026uuml;rttemberg\u003c/em\u003e, 35: 221\u0026ndash;304.\u003c/li\u003e\n\u003cli\u003eQuenstedt, F.A. (1884). \u003cem\u003eBegleitworte zur geognostischen Specialkarte von W\u0026uuml;rttemberg.\u003c/em\u003e\u003c/li\u003e\n\u003cli\u003e\u003cem\u003eAtlasbl\u0026auml;tter Ehingen, Biberach, Laupheim, Ochsenhausen\u003c/em\u003e. A. Kleeblatt \u0026amp; Comp.\u003c/li\u003e\n\u003cli\u003eQuenstedt, F.A. (1884). \u003cem\u003ePetrefactenkunde Deutschlands. 7. Band Gasteropoden\u003c/em\u003e. 867 pp. Leipzig (Fue`s).\u003c/li\u003e\n\u003cli\u003eRasser, M.W., Bechly, G., B\u0026ouml;ttcher, R., Ebner, M., Heizmann, E.P.J., H\u0026ouml;ltke, O., Joachim, C., Kern, A.K., Kovar-Eder, J., Nebelsick, J.H., Roth-Nebelsick, A., Schweigert, G., Schoch, R.R., Ziegler, R. (2013). The Randeck Maar: Palaeoenvironment and habitat differentiation of a Miocene lacustrine system. \u003cem\u003ePalaeogeography, Palaeoclimatology, Palaeoecology,\u003c/em\u003e 392: 426\u0026ndash; 453. http://dx.doi.org/10.1016/j.palaeo.2013.09.025\u003c/li\u003e\n\u003cli\u003eRollier, L. (1900). Vorl\u0026auml;ufige Notiz \u0026uuml;ber das Alter des Sylvanakalkes. \u003cem\u003eCentralblatt f\u0026uuml;r Mineralogie, Geologie und Palaeontologie\u003c/em\u003e, 1900: 89\u0026ndash;91.\u003c/li\u003e\n\u003cli\u003eRollier, L. (1903). \u0026Uuml;ber Diskordanzen im Schw\u0026auml;bischen Terti\u0026auml;r. \u003cem\u003eVierteljahrsschrift\u003c/em\u003e \u003cem\u003eder Naturforschenden Gesellschaft Z\u0026uuml;rich\u003c/em\u003e, 48: 307\u0026ndash;320.\u003c/li\u003e\n\u003cli\u003eSach, J. (1999). Litho- und biostratigraphische Untersuchungen in der Oberen S\u0026uuml;\u0026szlig;wassermolasse des Landkreises Biberach a. d. Ri\u0026szlig; (Oberschwaben).- \u003cem\u003eStuttgarter Beitr\u0026auml;ge zur Naturkunde\u003c/em\u003e, Serie B, 276:1-167.\u003c/li\u003e\n\u003cli\u003eSalvador, R.B. (2013a). The fossil land and freshwater mollusks of Sandelzhausen (Early/Middle Miocene, Germany): Caenogastropoda, Neritimorpha, lower Heterobranchia and Bivalvia. \u003cem\u003eStrombus\u003c/em\u003e, 20(1-2): 19\u0026ndash;26.\u003c/li\u003e\n\u003cli\u003eSalvador, R.B. (2013b). The fossil pulmonate snails of Sandelzhausen (Early/Middle Miocene, Germany): Succineidae, Testacelloidea and Helicoidea. \u003cem\u003eZootaxa\u003c/em\u003e, 3721(2): 157\u0026ndash;171. http://dx.doi.org/10.11646/zootaxa.3721.2.3\u003c/li\u003e\n\u003cli\u003eSalvador, R.B. (2014). The fossil land and freshwater snails of G\u0026uuml;ndlkofen (Middle Miocene, Germany). \u003cem\u003eZootaxa\u003c/em\u003e, 3785(2): 271\u0026ndash;287. https://doi.org/10.11646/zootaxa.3785.2.9\u003c/li\u003e\n\u003cli\u003eSalvador, R.B. (2015). The fossil pulmonate snails of Sandelzhausen (Early/Middle Miocene, Germany): Ellobiidae, Pupilloidea, and Clausilioidea. \u003cem\u003ePal\u0026auml;ontologische Zeitschrift\u003c/em\u003e, 89(1): 37\u0026ndash;50. http://dx.doi.org/10.1007/s12542-013-0210-4\u003c/li\u003e\n\u003cli\u003eSalvador, R.B. \u0026amp; Rasser, M.W. (2014). The fossil pulmonate snails of Sandelzhausen (Early/Middle Miocene, Germany) (Hygrophila, Punctoidea and limacoids). \u003cem\u003eArchiv fur Molluskenkunde,\u003c/em\u003e 143: 187\u0026ndash;202. http://dx.doi.org/10.1127/arch.moll/1869-0963/143/187-202\u003c/li\u003e\n\u003cli\u003eSalvador, R.B. \u0026amp; Rasser, M.W. (2016a). Fossil gastropods from the Middle Miocene of Bechingen and Daugendorf, southwestern Germany. \u003cem\u003eArchiv fur Molluskenkunde\u003c/em\u003e, 145(1): 111\u0026ndash; 124. DOI: 10.1127/arch.moll/1869-0963/145/111-124\u003c/li\u003e\n\u003cli\u003eSalvador, R.B. \u0026amp; Rasser, M.W. (2016b). The fossil land and freshwater snails of Oggenhausen (Middle Miocene, Germany). \u003cem\u003eRevista Brasileira de Paleontologia\u003c/em\u003e, 19(1): 41\u0026ndash;52. http://dx.doi.org/10.4072/rbp.2016.1.04\u003c/li\u003e\n\u003cli\u003eSalvador, R.B., H\u0026ouml;ltke, O. \u0026amp; Rasser, M.W. (2017). Fossil land and freshwater gastropods from the Miocene of Hohenmemmingen, Germany. \u003cem\u003ePalaeodiversity\u003c/em\u003e, 10: 41\u0026ndash;48. http://dx.doi.org/10.18476/pale.v10.a4\u003c/li\u003e\n\u003cli\u003eSalvador, R.B. \u0026amp; H\u0026ouml;ltke, O. \u0026amp; Rasser, M.W. (2018). Miocene continental gastropods from Dischingen, Germany. \u003cem\u003ePalaeodiversity\u003c/em\u003e, 11(1):11\u0026ndash;19. http://dx.doi.org/10.18476/pale.11.a2\u003c/li\u003e\n\u003cli\u003eSalvador, R.B., H\u0026ouml;ltke, O., Rasser, M.W. \u0026amp; Kadolsky, D. (2016a). Annotated type catalogue of the continental fossil gastropods in the Staatliches Museum f\u0026uuml;r Naturkunde Stuttgart, Germany. \u003cem\u003ePalaeodiversity\u003c/em\u003e, 9: 15\u0026ndash;70. http://dx.doi.org/10.18476/pale.v9.a3\u003c/li\u003e\n\u003cli\u003eSalvador, R.B., Prieto, J., Mayr, C., Rasser, M.W. (2016b). New gastropod assemblages from the Early/Middle Miocene of Riedensheim and Adelschlag-Fasanerie, southern Germany. \u003cem\u003eNeues Jahrbuch f\u0026uuml;r Geologie und Pal\u0026auml;ontologie, Abhandlungen\u003c/em\u003e, 279: 127\u0026ndash;154. http://dx.doi.org/10.1127/njgpa/2016/0546\u003c/li\u003e\n\u003cli\u003eSalvador, R.B., Rasser, M.W. \u0026amp; H\u0026ouml;ltke, O. (2015a). Fossil gastropods from Miocene Lake Randeck Maar and its hinterland (SW Germany). \u003cem\u003eNeues Jahrbuch f\u0026uuml;r Geologie und Pal\u0026auml;ontologie, Abhandlungen\u003c/em\u003e, 277(3): 251\u0026ndash;273. http://dx.doi.org/10.1127/njgpa/2015/0505\u003c/li\u003e\n\u003cli\u003eSalvador, R.B, Sach, V. \u0026amp; Valentas-Romera, B.L. (2015b). The fossil continental mollusks in the Upper Freshwater Molasse (Middle Miocene) of the districts of Biberach, Ravensburg and Neu-Ulm, Germany. \u003cem\u003eRevista Brasileira de Paleontologia\u003c/em\u003e, 18(2): 201\u0026ndash;216. http://dx.doi.org/10.4072/rbp.2015.2.02\u003c/li\u003e\n\u003cli\u003eSandberger, F. (1870\u0026ndash;1875). \u003cem\u003eDie Land- und S\u0026uuml;\u0026szlig;wasserconchylien der Vorwelt\u003c/em\u003e: livr. 1: 1\u0026ndash; 48, pl. 1\u0026ndash; 4, 1870; livr. 2\u0026ndash; 3: 49\u0026ndash; 96, pl. 5\u0026ndash; 12, 1870; livr. 4\u0026ndash; 5: 97\u0026ndash; 160, pl. 13\u0026ndash; 20, 1871; livr. 6\u0026ndash; 8: 161\u0026ndash; 256, pl. 21\u0026ndash; 32, 1872; livr. 9\u0026ndash; 10: 257\u0026ndash; 352, pl. 33\u0026ndash; 36, 1873; livr. 11: 353\u0026ndash; 616, 1875; livr. 12: 617\u0026ndash; 1000, 1875. Kreidel.\u003c/li\u003e\n\u003cli\u003eSchad, J. (1908). Beitrag zur Kenntnis des Terti\u0026auml;rs am Landgericht und Hochstr\u0026auml;\u0026szlig;. \u003cem\u003eJahreshefte des Vereins f\u0026uuml;r vaterl\u0026auml;ndische Naturkunde in W\u0026uuml;rttemberg\u003c/em\u003e, 64: 249\u0026ndash;304.\u003c/li\u003e\n\u003cli\u003eSchlickum, W.R. (1976). Die in der pleistoz\u0026auml;nen Gemeindekiesgrube von Zwiefaltendorf a. d. Donau abgelagerte Molluskenfauna der Silvanaschichten. Archiv f\u0026uuml;r Molluskenkunde 107(1/3): 1\u0026ndash;31.\u003c/li\u003e\n\u003cli\u003eSchlotheim, E.F. (1820). Die Petrefaktenkunde auf ihrem jetzigen Standpunkt durch die Beschreibung seiner Sammlung versteinerter und fossiler \u0026Uuml;berreste des Thier- und Pflanzenreichs der Vorwelt erl\u0026auml;utertBecker-Verlag.\u003c/li\u003e\n\u003cli\u003eSchwarz, F. (1913). \u003cem\u003eBeschreibung des Terti\u0026auml;rs im Tautschbuch-Emerberggebiet\u003c/em\u003e. 54 pp. Inaugural-Dissertation, T\u0026uuml;bingen (Laupp).\u003c/li\u003e\n\u003cli\u003eSchweigert, G. (1996). Vergleichende Faziesanalyse, Pal\u0026auml;o\u0026ouml;kologie und pal\u0026ouml;ogographisches Umfeld terti\u0026auml;rer S\u0026uuml;\u0026szlig;wasserkarbonate auf der westlichen Schw\u0026auml;bischen Alb und im Hegau (Baden-W\u0026uuml;rttemberg). \u003cem\u003eProfil\u003c/em\u003e, 9: 1\u0026ndash;100.\u003c/li\u003e\n\u003cli\u003eSimpson, G.G. (1943). Mammals and the nature of continents. \u003cem\u003eAmerican Journal of Science\u003c/em\u003e, 241: 1\u0026ndash;31. http://dx.doi.org/10.2475/ajs.241.1.1\u003c/li\u003e\n\u003cli\u003eSimpson, G.G. (1960). Notes on the measurement of faunal resemblance. \u003cem\u003eAmerican Journal of Science\u003c/em\u003e, 258A: 300\u0026ndash;311.\u003c/li\u003e\n\u003cli\u003eS\u0026oslash;rensen, T. (1948). A method of establishing groups of equal amplitude in plant sociology based on similarity of species and its application to analyses of the vegetation on Danish commons. \u003cem\u003eKongelige Danske Videnskabernes Selskab\u003c/em\u003e, 5(4): 1\u0026ndash;34.\u003c/li\u003e\n\u003cli\u003eWedel, J. (2008). Pleistocene molluscs from research boreholes in the Heidelberg Basin. \u003cem\u003eQuaternary Science Journal\u003c/em\u003e, 57(3\u0026ndash;4): 382\u0026ndash;402. http://dx.doi.org/10.3285/eg.57.3-4.6\u003c/li\u003e\n\u003cli\u003eWenz, W. (1920). \u0026Uuml;ber das Vorkommen von \u003cem\u003eCepaea eversa larteti\u003c/em\u003e (Boissy) in den schw\u0026auml;bischen Silvanaschichten und seine Bedeutung f\u0026uuml;r deren Gliederung. \u003cem\u003eSenckenbergiana,\u003c/em\u003e 2:151\u0026ndash;158.\u003c/li\u003e\n\u003cli\u003eWenz, W. (1921). Zur Frage der Altersstellung des schw\u0026auml;bischen Terti\u0026auml;rs. \u003cem\u003eCentralblatt f\u0026uuml;r Mineralogie, Geologie und Palaeontologie,\u003c/em\u003e 1921:559\u0026ndash;564.\u003c/li\u003e\n\u003cli\u003eWenz, W. (1923-1930). \u003cem\u003eGastropoda Extramarina Tertiaria. I-X\u003c/em\u003e. W. Junk.\u003c/li\u003e\n\u003cli\u003eWenz, W. (1924). Die Flammenmergel der Silvanaschichten und ihre Fauna. \u003cem\u003eJahresberichte und Mitteilungen des Oberrheinischen geologischen Vereins\u003c/em\u003e, N.F., 13: 181\u0026ndash;186.\u003c/li\u003e\n\u003cli\u003eWenz, W. (1933). Zur Land- und S\u0026uuml;\u0026szlig;wassermolluskenfauna der subalpinen Molasse des Pf\u0026auml;ndergebietes. \u003cem\u003eSenckenbergiana\u003c/em\u003e, 15: 7\u0026ndash;12.\u003c/li\u003e\n\u003cli\u003eWenz, W. (1935). Weitere Beitr\u0026auml;ge zur Land- und S\u0026uuml;\u0026szlig;wasser-Molluskenfauna der subalpinen Molasse des Pf\u0026auml;ndergebietes. \u003cem\u003eSenckenbergiana\u003c/em\u003e, 17: 223\u0026ndash;226.\u003c/li\u003e\n\u003cli\u003eWerner, W. (2014). \u003cem\u003eGauinger, Sonderbucher und Riedlinger Travertin\u003c/em\u003e. In: Werner, W., Wittenbrink, J., Bock, H. \u0026amp; Kimmig, B. (Eds.), \u003cem\u003eNaturwerksteine aus Baden-W\u0026uuml;rttemberg. Vorkommen, Beschaffenheit und Nutzung\u003c/em\u003e (pp. 279\u0026ndash;294). Landesamt f\u0026uuml;r Geologie, Rohstoffe und Bergbau.\u003c/li\u003e\n\u003cli\u003eZittel, K.A. (1885). \u003cem\u003eHandbuch der Palaeontologie. 1. Abteilung Palaeozoologie. II Band. Mollussca und Arthropoda\u003c/em\u003e. Oldenbourg.\u003c/li\u003e\n\u003cli\u003eZ\u0026ouml;belein, H.K. (1977). Anstehende und eiszeitlich verschleppte Obere S\u0026uuml;\u0026szlig;wasser-Molasse (Ober-Mioz\u0026auml;n) im Bereich des Riedlinger Beckens (Baden-W\u0026uuml;rttemberg). \u003cem\u003eMitteilungen der Bayerischen Staatssammlung f\u0026uuml;r Pal\u0026auml;ontologie und Historische Geologie\u003c/em\u003e, 17: 291\u0026ndash;334.\u003c/li\u003e\n\u003c/ol\u003e"},{"header":"Table 1","content":"\u003cp\u003eTable 1. Deposits and references.\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.53301886792453%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eLocality\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.320754716981131%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eInside NAFB?\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.61320754716981%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eCountry\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.495283018867925%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eNr. Species\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"41.0377358490566%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eReference(s)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.53301886792453%\" valign=\"top\"\u003e\n \u003cp\u003eAdelschlag-Fasanerie\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.320754716981131%\" valign=\"top\"\u003e\n \u003cp\u003eyes\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.61320754716981%\" valign=\"top\"\u003e\n \u003cp\u003eGermany (BY)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.495283018867925%\" valign=\"top\"\u003e\n \u003cp\u003e8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"41.0377358490566%\" valign=\"top\"\u003e\n \u003cp\u003eSalvador et al. (2016b)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.53301886792453%\" valign=\"top\"\u003e\n \u003cp\u003eAltheim near Ehingen/Donau\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.320754716981131%\" valign=\"top\"\u003e\n \u003cp\u003eyes\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.61320754716981%\" valign=\"top\"\u003e\n \u003cp\u003eGermany (BW)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.495283018867925%\" valign=\"top\"\u003e\n \u003cp\u003e13\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"41.0377358490566%\" valign=\"top\"\u003e\n \u003cp\u003eH\u0026ouml;ltke et al. (2018)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.53301886792453%\" valign=\"top\"\u003e\n \u003cp\u003eAndelfingen\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.320754716981131%\" valign=\"top\"\u003e\n \u003cp\u003eyes\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.61320754716981%\" valign=\"top\"\u003e\n \u003cp\u003eGermany (BW)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.495283018867925%\" valign=\"top\"\u003e\n \u003cp\u003e8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"41.0377358490566%\" valign=\"top\"\u003e\n \u003cp\u003eH\u0026ouml;ltke et al. (2018)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.53301886792453%\" valign=\"top\"\u003e\n \u003cp\u003eBaach\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.320754716981131%\" valign=\"top\"\u003e\n \u003cp\u003eyes\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.61320754716981%\" valign=\"top\"\u003e\n \u003cp\u003eGermany (BW)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.495283018867925%\" valign=\"top\"\u003e\n \u003cp\u003e7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"41.0377358490566%\" valign=\"top\"\u003e\n \u003cp\u003eH\u0026ouml;ltke et al. (2018)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.53301886792453%\" valign=\"top\"\u003e\n \u003cp\u003eBechingen\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.320754716981131%\" valign=\"top\"\u003e\n \u003cp\u003eyes\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.61320754716981%\" valign=\"top\"\u003e\n \u003cp\u003eGermany (BW)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.495283018867925%\" valign=\"top\"\u003e\n \u003cp\u003e17\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"41.0377358490566%\" valign=\"top\"\u003e\n \u003cp\u003eSalvador \u0026amp; Rasser (2016a)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.53301886792453%\" valign=\"top\"\u003e\n \u003cp\u003eD\u0026auml;chingen\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.320754716981131%\" valign=\"top\"\u003e\n \u003cp\u003eyes\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.61320754716981%\" valign=\"top\"\u003e\n \u003cp\u003eGermany (BW)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.495283018867925%\" valign=\"top\"\u003e\n \u003cp\u003e6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"41.0377358490566%\" valign=\"top\"\u003e\n \u003cp\u003eH\u0026ouml;ltke et al. (2018)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.53301886792453%\" valign=\"top\"\u003e\n \u003cp\u003eDaugendorf\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.320754716981131%\" valign=\"top\"\u003e\n \u003cp\u003eyes\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.61320754716981%\" valign=\"top\"\u003e\n \u003cp\u003eGermany (BW)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.495283018867925%\" valign=\"top\"\u003e\n \u003cp\u003e7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"41.0377358490566%\" valign=\"top\"\u003e\n \u003cp\u003eSalvador \u0026amp; Rasser (2016)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.53301886792453%\" valign=\"top\"\u003e\n \u003cp\u003eDischingen\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.320754716981131%\" valign=\"top\"\u003e\n \u003cp\u003eyes\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.61320754716981%\" valign=\"top\"\u003e\n \u003cp\u003eGermany (BW)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.495283018867925%\" valign=\"top\"\u003e\n \u003cp\u003e25\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"41.0377358490566%\" valign=\"top\"\u003e\n \u003cp\u003eSalvador et al. 2018\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.53301886792453%\" valign=\"top\"\u003e\n \u003cp\u003eEhingen/Donau\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.320754716981131%\" valign=\"top\"\u003e\n \u003cp\u003eyes\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.61320754716981%\" valign=\"top\"\u003e\n \u003cp\u003eGermany (BW)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.495283018867925%\" valign=\"top\"\u003e\n \u003cp\u003e6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"41.0377358490566%\" valign=\"top\"\u003e\n \u003cp\u003eH\u0026ouml;ltke et al. (2018)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.53301886792453%\" valign=\"top\"\u003e\n \u003cp\u003eEmerberg\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.320754716981131%\" valign=\"top\"\u003e\n \u003cp\u003eyes\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.61320754716981%\" valign=\"top\"\u003e\n \u003cp\u003eGermany (BW)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.495283018867925%\" valign=\"top\"\u003e\n \u003cp\u003e18\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"41.0377358490566%\" valign=\"top\"\u003e\n \u003cp\u003eH\u0026ouml;ltke et al. (2018)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.53301886792453%\" valign=\"top\"\u003e\n \u003cp\u003eFriedingen\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.320754716981131%\" valign=\"top\"\u003e\n \u003cp\u003eyes\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.61320754716981%\" valign=\"top\"\u003e\n \u003cp\u003eGermany (BW)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.495283018867925%\" valign=\"top\"\u003e\n \u003cp\u003e8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"41.0377358490566%\" valign=\"top\"\u003e\n \u003cp\u003eWenz (1923)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.53301886792453%\" valign=\"top\"\u003e\n \u003cp\u003eGeorgensgm\u0026uuml;nd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.320754716981131%\" valign=\"top\"\u003e\n \u003cp\u003eyes\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.61320754716981%\" valign=\"top\"\u003e\n \u003cp\u003eGermany (BY)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.495283018867925%\" valign=\"top\"\u003e\n \u003cp\u003e7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"41.0377358490566%\" valign=\"top\"\u003e\n \u003cp\u003eBerger (2010)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.53301886792453%\" valign=\"top\"\u003e\n \u003cp\u003eG\u0026uuml;ndlkofen\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.320754716981131%\" valign=\"top\"\u003e\n \u003cp\u003eyes\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.61320754716981%\" valign=\"top\"\u003e\n \u003cp\u003eGermany (BY)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.495283018867925%\" valign=\"top\"\u003e\n \u003cp\u003e7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"41.0377358490566%\" valign=\"top\"\u003e\n \u003cp\u003eSalvador (2014)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.53301886792453%\" valign=\"top\"\u003e\n \u003cp\u003eHausen ob Allmendingen\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.320754716981131%\" valign=\"top\"\u003e\n \u003cp\u003eyes\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.61320754716981%\" valign=\"top\"\u003e\n \u003cp\u003eGermany (BW)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.495283018867925%\" valign=\"top\"\u003e\n \u003cp\u003e11\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"41.0377358490566%\" valign=\"top\"\u003e\n \u003cp\u003eH\u0026ouml;ltke et al. (2018)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.53301886792453%\" valign=\"top\"\u003e\n \u003cp\u003eHengen\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.320754716981131%\" valign=\"top\"\u003e\n \u003cp\u003eno\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.61320754716981%\" valign=\"top\"\u003e\n \u003cp\u003eGermany (BW)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.495283018867925%\" valign=\"top\"\u003e\n \u003cp\u003e11\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"41.0377358490566%\" valign=\"top\"\u003e\n \u003cp\u003eEhrat \u0026amp; Jooss (1921), collection material\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.53301886792453%\" valign=\"top\"\u003e\n \u003cp\u003eHohenmemmingen\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.320754716981131%\" valign=\"top\"\u003e\n \u003cp\u003eyes\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.61320754716981%\" valign=\"top\"\u003e\n \u003cp\u003eGermany (BW)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.495283018867925%\" valign=\"top\"\u003e\n \u003cp\u003e24\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"41.0377358490566%\" valign=\"top\"\u003e\n \u003cp\u003eSalvador et al. (2017)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.53301886792453%\" valign=\"top\"\u003e\n \u003cp\u003eM\u0026ouml;rsingen\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.320754716981131%\" valign=\"top\"\u003e\n \u003cp\u003eyes\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.61320754716981%\" valign=\"top\"\u003e\n \u003cp\u003eGermany (BW)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.495283018867925%\" valign=\"top\"\u003e\n \u003cp\u003e36\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"41.0377358490566%\" valign=\"top\"\u003e\n \u003cp\u003eSchlickum (1976), collection material\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.53301886792453%\" valign=\"top\"\u003e\n \u003cp\u003eMundingen\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.320754716981131%\" valign=\"top\"\u003e\n \u003cp\u003eyes\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.61320754716981%\" valign=\"top\"\u003e\n \u003cp\u003eGermany (BW)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.495283018867925%\" valign=\"top\"\u003e\n \u003cp\u003e6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"41.0377358490566%\" valign=\"top\"\u003e\n \u003cp\u003eWenz (1923)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.53301886792453%\" valign=\"top\"\u003e\n \u003cp\u003eOggenhausen\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.320754716981131%\" valign=\"top\"\u003e\n \u003cp\u003eyes\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.61320754716981%\" valign=\"top\"\u003e\n \u003cp\u003eGermany (BY)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.495283018867925%\" valign=\"top\"\u003e\n \u003cp\u003e8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"41.0377358490566%\" valign=\"top\"\u003e\n \u003cp\u003eSalvador \u0026amp; Rasser (2016b)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.53301886792453%\" valign=\"top\"\u003e\n \u003cp\u003ePf\u0026auml;nder near Bregenz\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.320754716981131%\" valign=\"top\"\u003e\n \u003cp\u003eyes\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.61320754716981%\" valign=\"top\"\u003e\n \u003cp\u003eAustria (Vorarlberg)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.495283018867925%\" valign=\"top\"\u003e\n \u003cp\u003e8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"41.0377358490566%\" valign=\"top\"\u003e\n \u003cp\u003eJooss (1910, 1915), Wenz (1933, 1935)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.53301886792453%\" valign=\"top\"\u003e\n \u003cp\u003eRandecker Maar\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.320754716981131%\" valign=\"top\"\u003e\n \u003cp\u003eno\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.61320754716981%\" valign=\"top\"\u003e\n \u003cp\u003eGermany (BW)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.495283018867925%\" valign=\"top\"\u003e\n \u003cp\u003e17\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"41.0377358490566%\" valign=\"top\"\u003e\n \u003cp\u003eSalvador et al. (2015a)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.53301886792453%\" valign=\"top\"\u003e\n \u003cp\u003eRennertshofen-Riedensheim\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.320754716981131%\" valign=\"top\"\u003e\n \u003cp\u003eyes\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.61320754716981%\" valign=\"top\"\u003e\n \u003cp\u003eGermany\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.495283018867925%\" valign=\"top\"\u003e\n \u003cp\u003e13\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"41.0377358490566%\" valign=\"top\"\u003e\n \u003cp\u003eSalvador et al. (2016b)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.53301886792453%\" valign=\"top\"\u003e\n \u003cp\u003eSandelzhausen\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.320754716981131%\" valign=\"top\"\u003e\n \u003cp\u003eyes\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.61320754716981%\" valign=\"top\"\u003e\n \u003cp\u003eGermany (BW)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.495283018867925%\" valign=\"top\"\u003e\n \u003cp\u003e14\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"41.0377358490566%\" valign=\"top\"\u003e\n \u003cp\u003eSalvador (2013a, 2013b, 2015); Salvador \u0026amp; Rasser (2014)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.53301886792453%\" valign=\"top\"\u003e\n \u003cp\u003eStoffelberg near Ehingen/Donau\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.320754716981131%\" valign=\"top\"\u003e\n \u003cp\u003eyes\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.61320754716981%\" valign=\"top\"\u003e\n \u003cp\u003eGermany (BW)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.495283018867925%\" valign=\"top\"\u003e\n \u003cp\u003e14\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"41.0377358490566%\" valign=\"top\"\u003e\n \u003cp\u003eH\u0026ouml;ltke et al. (2018)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.53301886792453%\" valign=\"top\"\u003e\n \u003cp\u003eUndorf\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.320754716981131%\" valign=\"top\"\u003e\n \u003cp\u003eyes\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.61320754716981%\" valign=\"top\"\u003e\n \u003cp\u003eGermany (BY)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.495283018867925%\" valign=\"top\"\u003e\n \u003cp\u003e28\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"41.0377358490566%\" valign=\"top\"\u003e\n \u003cp\u003eClessin (1911)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.53301886792453%\" valign=\"top\"\u003e\n \u003cp\u003eVermes\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.320754716981131%\" valign=\"top\"\u003e\n \u003cp\u003eyes\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.61320754716981%\" valign=\"top\"\u003e\n \u003cp\u003eSwitzerland (Jura)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.495283018867925%\" valign=\"top\"\u003e\n \u003cp\u003e8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"41.0377358490566%\" valign=\"top\"\u003e\n \u003cp\u003eWenz (1923)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.53301886792453%\" valign=\"top\"\u003e\n \u003cp\u003eZwiefalten\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.320754716981131%\" valign=\"top\"\u003e\n \u003cp\u003eyes\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.61320754716981%\" valign=\"top\"\u003e\n \u003cp\u003eGermany (BW)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.495283018867925%\" valign=\"top\"\u003e\n \u003cp\u003e17\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"41.0377358490566%\" valign=\"top\"\u003e\n \u003cp\u003eH\u0026ouml;ltke et al. (2018)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.53301886792453%\" valign=\"top\"\u003e\n \u003cp\u003eZwiefaltendorf\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.320754716981131%\" valign=\"top\"\u003e\n \u003cp\u003eyes\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.61320754716981%\" valign=\"top\"\u003e\n \u003cp\u003eGermany (BW)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.495283018867925%\" valign=\"top\"\u003e\n \u003cp\u003e36\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"41.0377358490566%\" valign=\"top\"\u003e\n \u003cp\u003eSchlickum (1976), H\u0026ouml;ltke et al. (2016)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"cluster analysis, Gastropoda, similarity indices, Upper Freshwater Molasse","lastPublishedDoi":"10.21203/rs.3.rs-2752635/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-2752635/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThe \u003cem\u003esilvana\u003c/em\u003e-beds are the main component of the Upper Freshwater Molasse (\u0026ldquo;Obere S\u0026uuml;\u0026szlig;wassermolasse\u0026rdquo; or OSM) in southern Germany, Austria and Switzerland, named after the land snail \u003cem\u003ePalaeotachea silvana\u003c/em\u003e (Klein, \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e1853\u003c/span\u003e). After initial disputes in the literature, in the 1920\u0026rsquo;s, malacologist Wilhelm Wenz better defined the \u003cem\u003esilvana\u003c/em\u003e-beds lithology and its three informal subunits based on terrestrial gastropods. The \u003cem\u003esilvana\u003c/em\u003e-beds mainly consist of limestone, marl and clay, eventually with sandy components. Despite remaining an informal stratigraphical unit, the \u003cem\u003esilvana\u003c/em\u003e-beds is a practical tool for stratigraphic determination in the field and for paleomalacology studies. Information about the land snail communities of the \u003cem\u003esilvana\u003c/em\u003e-beds are scattered in the literature and museum collections. Therefore, here we compile all this information and conduct a detailed biogeographic comparison of these localities\u0026rsquo; snail faunas, based on similarity indices. For that purpose, all communities (containing at least six terrestrial gastropod species, from a total of 28 localities) within the \u003cem\u003esilvana\u003c/em\u003e-beds and the adjoining coeval Bad Urach volcanic region were analyzed, based on four similarity indices and follow-up cluster analyses. The land snail communities of the \u003cem\u003esilvana\u003c/em\u003e-beds formed six clusters (defined by ca. 50% similarity) according to the S\u0026oslash;rensen-Dice index, five clusters by the Kulczynski and Ochiai indices, and four ones by the Simpson index. Despite some differences in the resulting dendrograms, enough similarity in the clustering was found for a meaningful comparison. The resulting clusters can be largely explained by geographic position, as is the case of the two main groupings recovered, representing the German Bavarian and Baden-W\u0026uuml;rttemberg states. Some inconsistencies in the composition of the clusters might be explained by different stratigraphic positions within the \u003cem\u003esilvana\u003c/em\u003e-beds, distinct paleoenvironmental conditions influencing the faunal composition, or the absence of key species.\u003c/p\u003e","manuscriptTitle":"Paleobiogeography of the land snail communities of the Middle Miocene silvana-beds of Germany","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2023-04-25 13:09:30","doi":"10.21203/rs.3.rs-2752635/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"307b531b-0069-43f2-8c8b-0318c54eb79e","owner":[],"postedDate":"April 25th, 2023","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2023-06-27T08:32:04+00:00","versionOfRecord":[],"versionCreatedAt":"2023-04-25 13:09:30","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-2752635","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-2752635","identity":"rs-2752635","version":["v1"]},"buildId":"7rjqhiLT3MXkJMwkYKINL","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}