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Snake Fauna (Reptilia: Serpentes) from the Early/Middle Miocene of Sandelzhausen and Rothenstein 13 (Germany)

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Snake Fauna (Reptilia: Serpentes) from the Early/Middle Miocene of Sandelzhausen and Rothenstein 13 (Germany) View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by RERO DOC Digital Library Pala¨ontol Z (2009) 83:55–66 DOI 10.1007/s12542-009-0009-5 RESEARCH PAPER Snake fauna (Reptilia: Serpentes) from the Early/Middle Miocene of Sandelzhausen and Rothenstein 13 (Germany) Zbigniew Szyndlar Received: 21 September 2006 / Accepted: 26 April 2007 / Published online: 3 February 2009 Ó Springer-Verlag 2009 Abstract The Early/Middle Miocene (European Land Faunen repra¨sentieren ein U¨ bergangsstadium zwischen Mammal Zone MN5) localities of Sandelzhausen and Ro- Vergesellschaftungen anderer mittel- und westeuropa¨ischer thenstein 13 in southern Germany have yielded remains of Fundstellen aus dem spa¨ten Unter- und Mittelmioza¨n. Die about 13 ophidian taxa: Eoanilius sp. (Aniliidae), Bavar- Schlangenfossilien zeigen fu¨r Sandelzhausen und Rothen- ioboa ultima (Boidae), ‘‘Coluber’’ sp., ?Telescopus sp., stein13 ein warmes, aber nicht tropisches oder Natrix sp., cf. Natrix sp., cf. ‘‘Neonatrix’’ sp., unidentified subtropisches Klima an. ‘‘colubrines’’ and ‘‘natricines’’ (Colubridae), Naja sp., an unidentified elapid (Elapidae), Vipera sp. (‘‘Oriental Schlu¨sselwo¨rter Reptilia Á Serpentes Á Mioza¨n Á viper’’), Vipera sp. (‘‘aspis complex’’) (Viperidae). Both Deutschland Á Taxonomische Beschreibung Á faunas document a transitional phase from those reported Faunenzusammensetzung Á Pala¨oo¨kologie from several late Early and Middle Miocene sites of Cen- tral and Western Europe. The climates of Sandelzhausen and Rothenstein 13, as indicated by ophidian fossils, were Introduction warm, although not tropical or subtropical. The end of the Early Miocene (European Land Mammal Keywords Reptilia Á Serpentes Á Miocene Á Germany Á Zone MN4), when ‘‘archaic’’ snakes were replaced by their Taxonomic description Á Faunal composition Á ‘‘modern’’ successors, was a turning point in the history of Palaeoecology the European ophidian faunas (Szyndlar and Schleich 1993; Szyndlar and Rage 2003; Rage and Szyndlar 2005). Kurzfassung Aus den unter-/mittelmioza¨nen (Europa¨- The composition of snake assemblages representing MN4 ische Landsa¨uger Zone MN5) Fossilfundstellen has been fairly well studied in the last decade, but those Sandelzhausen und Rothenstein 13 in Su¨ddeutschland sind inhabiting Europe in the latest Early/early Middle Miocene Reste von 13 Schlangentaxa u¨berliefert: Eoanilius sp. (MN5) have not attracted much attention to date. Thus far, (Aniliidae), Bavarioboa ultima (Boidae), ‘‘Coluber’’ sp., most of our knowledge of the MN5 snakes has been ?Telescopus sp., Natrix sp., cf. Natrix sp., cf. ‘‘Neonatrix’’ derived from extrapolations of the content of the old levels sp., nicht na¨her identifizierte ‘‘colubrine’’ und ‘‘natricine’’ of the French locality La Grive (MN7 ? 8) (e.g. Szyndlar Schlangen (Colubridae), Naja sp., ein nicht na¨her identi- and Rage 2003). In this context, the fossil remains of fizierter Vertreter der Elapidae, Vipera sp. (‘‘Orientalische Sandelzhausen representing the European Land Mammal Viper’’), Vipera sp. (‘‘aspis-Komplex’’) (Viperidae). Beide Zone MN5 (Heissig 1997;Ro¨ssner 1997; Fahlbusch 2003; Hokkanen and Evans 2006; Moser et al. 2009 this volume) are of special importance for the knowledge of the history Z. Szyndlar (&) of European ophidian faunas. Although the preservation of Institute of Systematics and Evolution of Animals, the specimens is too poor to provide detailed information, Polish Academy of Sciences, Slawkowska 17, 31-016 Krako´w, Poland the snake fossils found in both sites give evidence for the e-mail: [email protected] faunal turnover in European snakes. 123 56 Z. Szyndlar The snake assemblage from Sandelzhausen is presented following snakes: Bavarioboa ultima (family Boidae); here along with that from Rothenstein 13 because of close Natrix sp. and cf. ‘‘Neonatrix’’ sp. (family Colubridae, taxonomical similarities at genus level (unfortunately, most subfamily ‘‘Natricinae’’); Naja sp. (family Elapidae); Vi- fossil remains cannot be identified to the species level) in pera sp. (‘‘aspis complex’’) (family Viperidae). In addition, both faunas. Rothenstein 13 is a karstic site located at the nine fragmentary vertebrae were identified to the (sub-) Franconian Alb in southern Germany north of the Molasse family level only (‘‘Colubrinae’’ indet. and Elapidae in- Basin (488970N, 118060E) with micromammal remains det.); 64 bony elements were not identified. indicating a biostratigraphic age of MN5 (E.P.J. Heizmann, It is worth adding that, in the case of a few snake taxa personal communication). So far, several genera of from Sandelzhausen, a striking disproportion in the number amphibians and reptiles found in the latter locality have of various skeletal elements can be observed. For instance, been reported (Bo¨hme and Ilg 2003; Szyndlar and Schleich the ophidian material contains more than 50 isolated ven- 1993; Szyndlar and Rage 2003). omous fangs of adult individuals of Vipera sp., but there Up to now, except for Bavarioboa ultima (Szyndlar and are only a dozen vertebrae referred to the same genus. It is Rage 2003), snake remains from the aforementioned clear that 50 fangs must have belonged to 25–50 individ- localities have not been described. The presence of a few uals, so it is surprising that the fangs are accompanied by ophidian taxa in both sites was mentioned by Szyndlar and so few vertebrae. It is difficult to hypothesise what mech- Schleich (1993) and Szyndlar and Bo¨hme (1993) (at that anisms may have been behind this disproportionate number time estimated to be of MN6 age). Szyndlar and Rage of some fossil remains. It seems, however, that the dis- (2003) listed the snake fauna of Rothenstein 13. proportions may have resulted from the applied excavation procedure rather than from the influence of taphonomic processes. The anatomical terminology used is according to Materials and methods Szyndlar and Rage (2003); see Fig. 1 for details. The entire collection of fossil snake material from San- delzhausen available for study belongs to the collection of Systematic palaeontology the Bayerische Staatssammlung fu¨r Pala¨ontologie und Geologie in Munich (BSPG). Most of the fossils used in the Class Reptilia Laurenti, 1768. present study, loaned by Prof. Volker Fahlbusch and Dr. Order Squamata Oppel, 1811. Gertrud Ro¨ssner in September 2003, are vertebral frag- Suborder Serpentes Linnaeus, 1758. ments. Due to the poor state of preservation, identification Infraorder Alethinophidia Nopcsa, 1923. of the material was impossible below the generic level. Of Family Aniliidae Fitzinger, 1826. 346 ophidian fossils available for study, 145 vertebrae and Genus Eoanilius (Rage 1974). venomous fangs were identified as belonging to the fol- Type species: Eoanilius europae (Rage 1974). lowing genera: Eoanilius sp. (family Aniliidae), ‘‘Coluber’’ Eoanilius sp. sp. and ?Telescopus sp. (family Colubridae, subfamily Figure 1. ‘‘Colubrinae’’), Natrix sp., cf. Natrix sp. and cf. ‘‘Neona- Material: trix’’ sp. (family Colubridae, subfamily ‘‘Natricinae’’); Ten trunk vertebrae from Sandelzhausen. Naja sp. (family Elapidae); Vipera sp. (‘‘Oriental viper’’) Description: (family Viperidae). A further 67 fragmentary vertebrae and The vertebrae are characterized by very small dimen- 4 fragments of toothed bones were identified to the (sub-) sions and relatively simple morphology. In the largest family level only (‘‘Colubrinae’’ indet., ‘‘Natricinae’’ in- (incomplete) vertebra the centrum length is 2.6 mm; in det., Elapidae indet.); 130 caudal vertebrae were not remaining vertebrae it ranges between 1.8 and 2.1 mm. identified. Most vertebrae come from the middle trunk portion of In addition to the above, 48 ophidian fragmentary trunk the vertebral column. The centrum is as wide as long, and vertebrae from Sandelzhausen were studied by Szyndlar cylindrical. The haemal keel is very indistinct (though and Schleich at the end of 1991 and mentioned in their broad), accompanied by very shallow subcentral grooves. joint paper published in 1993. The remains were identified The neural arch is strongly depressed. The neural spine is as Coluber sp., Naja sp., and Vipera sp. (‘‘Oriental viper’’). extremely low; it occupies one-third the neural arch length. The material from Rothenstein 13 belongs to the collection The pre- and postzygapophyseal articular facets are of the Staatliches Museum fu¨r Naturkunde in Stuttgart relatively large, oval-shaped and elongated. The prezyga- (SMNS). It consists of 143 (mainly fragmentary) vertebrae pophyseal processes are very short and hardly visible in and 3 fragments of cranial bones. Of them, 73 vertebrae dorsal view. The zygosphenal roof is either three-lobed in and 1 compound were identified as belonging to the dorsal view or is devoid of the central lobe. The cotyle and 123 Snake fauna from the Early/Middle Miocene of Sandelzhausen and Rothenstein 13 57 Fig. 1 Two trunk vertebrae (a–e BSPG 1959 II 15967; f–h BSPG prezygapophyseal articular surface; prp: prezygapophyseal process; 1959 II 15966) of Eoanilius sp. from Sandelzhausen. cd: condyle; ct: sf: subcentral foramen; sg: subcentral groove; sr: subcentral ridge; z: cotyle; hk: haemal keel; lf: lateral foramen; na: neural arch; nc: neural zygosphene; zy: zygantrum. a, f left lateral views; b, g dorsal views; canal; ns: neural spine; pd: paradiapophysis; po: postzygapophysis; c, h ventral views; d anterior view; e posterior view poa: postzygapophyseal articular surface; pr: prezygapophysis; pra: condyle are suborbicular or
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