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

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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 westeuropaïscher 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 Serpentes Mioza¨n viper’’), Vipera sp. (‘‘aspis complex’’) (Viperidae). Both Deutschland Taxonomische Beschreibung faunas document a transitional phase from those reported Faunenzusammensetzung Palaöo¨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 Serpentes Miocene Germany Zone MN4), when ‘‘archaic’’ snakes were replaced by their Taxonomic description Faunal composition ‘‘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 Landsaüger 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 Palaöntologie 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 ﬂattened dorso-ventrally. The younger remains of this snake, usually referred to the paradiapophyses are relatively large, with no clear division species E. oligocenicus (Szyndlar 1994), were reported into para- and diapophyseal portions. The lateral and sub- from several German and French localities, dated from central foramina are distinct; the paracotylar foramina are the Early Oligocene to Early Miocene (see Szyndlar and absent. Rage 2003, and references therein). Additional remains The vertebrae interpreted as coming from the posterior of Eoanilius, not included in the list by Szyndlar and trunk portion of the column are distinctly longer than those Rage, were also found in two French Oligocene locali- from the middle trunk portion; they have more distinct ties, Lebratie`res 14 and La Colombie`re (Rage and haemal keels and deeper subcentral grooves. Szyndlar, unpublished), as well as in the Early Miocene Comments: (MN3 or 3–4) of Agramo´n in Spain (Szyndlar and Al- The vertebrae are clearly referable to the extinct Eo- fe´rez 2005). anilius, a member of the family Aniliidae, the present-day Interestingly, Eoanilius was a dominant element in distribution of which is restricted to tropical America. middle Early Miocene (MN2 and 3) snake assemblages of The oldest remains of the extinct genus Eoanilius, southern Germany (Szyndlar and Schleich 1993). The represented by the type-species E. europae (Rage 1974), vertebrae from Sandelzhausen are morphologically and come from the late Eocene of France (Rage 1974), metrically very similar to those of Eoanilius sp. reported Britain (Milner et al. 1982) and Spain (Szyndlar and from the locality of Petersbuch 2 (MN4) (Szyndlar and Alfe´rez 2005). Eoanilius is considered the only ophidian Schleich 1993). genus that survived the ‘‘Grande Coupure’’ events at the Sandelzhausen is the geologically youngest fossil site Eocene/Oligocene boundary in Europe. Geologically yielding remains of Eoanilius.

123 58 Z. Szyndlar

Family Boidae Gray, 1825. The centrum is triangular in ventral view and it is slightly Genus Bavarioboa Szyndlar and Schleich, 1993. longer than wide. The subcentral ridges are well developed. Type species: Bavarioboa hermi Szyndlar and Schleich, The haemal keel is distinct and cuneate-shaped. The neural 1993. arch is moderately vaulted, not accompanied by epizyga- Bavarioboa ultima Szyndlar and Rage, 2003. pophyseal spines. The neural spine (almost completely Material: preserved in one vertebra only; Fig. 2j) is approximately as Twenty-two trunk vertebrae, one cloacal vertebra, and high as long; the posterior margin of the spine slightly one caudal vertebra from Rothenstein 13. overhangs. The zygosphene (completely preserved in few Comments: vertebrae) is roughly straight. The prezygapophyseal Szyndlar and Schleich (1993), who erected the extinct articular facets are oval. The prezygapophyseal processes boid genus Bavarioboa, considered the snake from Ro- (preserved in three vertebrae) are either as long as the thenstein 13 a member of the type (then the only known) articular facet and obtuse (Fig. 2h) or somewhat shorter species, i.e. Bavarioboa hermi from Petersbuch 2. Szyndlar than the facet and acute (Fig. 2i). The paradiapophyses are and Rage (2003) described the remains from Rothen- moderately well developed, with the dia- and parapophy- stein 13 as a distinct species, named Bavarioboa ultima. seal portions of roughly equal length. The cotyle and They based their description on 18 trunk vertebrae. In condyle are slightly depressed dorso-ventrally. The lateral, addition, the ophidian material from Rothenstein 13 con- subcentral and paracotylar foramina are distinct. tains four other trunk vertebrae, one cloacal (sacral) Comments: vertebra and one caudal vertebra referable to this snake. In its vertebral morphology, the snake from Sandelz- See Szyndlar and Rage (2003) for detailed generic and hausen most approaches the conditions observed in the specific diagnoses of this snake. vertebrae of four large-sized (and perhaps closely related) Representatives of the ophidian superfamily Booidea fossil species, namely Coluber dolnicensis, C. caspioides, (mainly members of the family Boidae) were dominant C. suevicus and C. pouchetii. A single maxillary fragment, elements in snake faunas throughout the Paleogene, in owing to the presence of a completely preserved ‘‘Coluber- Europe and elsewhere. Members of the extinct genus Ba- like’’ ectopterygoid process, is also temporarily referred to varioboa, along with the erycine Bransateryx, were the the same snake. All the ophidian species mentioned in the most common European snakes in the Middle and Late preceding paragraph are referred to the informal subfamily Oligocene; Bavarioboa disappeared from Europe at the end ‘‘Colubrinae’’ (or ‘‘colubrines’’), characterized by the of the Oligocene, but it reappeared in the continent at the absence of hypapophyses in their middle trunk vertebrae. It end of the Early Miocene (Szyndlar and Rage 2003; Rage is worth adding that, in the recent past, the generic name and Szyndlar 2005). Bavarioboa ultima from Rothen- Coluber encompassed as many as nine species of living stein 13, along with Bavarioboa sp. from the French European ‘‘colubrines’’, today classified in the genera locality Isle d’Abeau, is the geologically youngest repre- Hemorrhois, Hierophis and Platyceps. The most likely sentative of the genus (Szyndlar and Rage 2003). Most allocation of the fossil group dolnicensis–caspioides–su- European boid snakes became extinct before the end of the evicus–pouchetii is in the genus Hierophis. In this paper I Middle Miocene (MN7 ? 8); the only representatives of use the traditional name ‘‘Coluber’’, mainly because this the family that survived until today in the southeastern part name is still in use in the European paleontological liter- of the European continent are small sand boas (genus Eryx). ature. However, I put the generic name in quotation marks. Family Colubridae Oppel, 1811. Rage and Bailon (2005) listed vertebral characteristics Subfamily ‘‘Colubrinae’’. enabling to distinguish remains of the above four fossil Genus Coluber (s.l.) Linnaeus, 1758. species. Unfortunately, regarding the snake from Sandelz- Type species: Coluber constrictor Linnaeus, 1758. hausen, the fragmentary nature of its vertebrae does not ‘‘Coluber’’ sp. enable to identify it to the specific level. Figure 2. The oldest fossil record of snakes belonging to the dis- Material: cussed group comes from the Czech locality of Merkur- Sixty-five trunk vertebrae and one maxillary fragment North (MN3) from which Ivanov (2002) reported the from Sandelzhausen. presence of three species of the complex (Coluber dol- Description: nicensis, C. caspioide , C. pouchetii) altogether. In the The discussed vertebrae, all preserved in more or less final Early and Middle Miocene these species became fragmentary state, come from the middle trunk portion of widespread in the European continent (e.g. Szyndlar and the column. They belonged to large-sized snakes; the Schleich 1993; Auge´ and Rage 2000; Ivanov 2000; Rage centrum length of the largest vertebra (Fig. 2a) is 9.7 mm. and Bailon 2005). The geologically youngest record is that The vertebrae are relatively high and short in lateral view. of cf. C. pouchetii from the early Late Miocene (MN9) of 123 Snake fauna from the Early/Middle Miocene of Sandelzhausen and Rothenstein 13 59

Fig. 2 Four trunk vertebrae (a BSPG 1959 II 8094; b–f BSPG 1952 II 16092; g, h BSPG 1959 II 16093; i, j BSPG 1959 II 16131) of Coluber sp. from Sandelzhausen. a, b, j left lateral views; c, h, i dorsal views; d anterior view; e, g ventral views; f posterior view the Hungarian locality of Rudaba´nya (Szyndlar 2005). The reported from several localities of eastern and central alleged closest living relative of the Miocene ‘‘Coluber’’ is Europe, ranging in age from the Early Miocene (MN4) to Hierophis (formerly Coluber) caspius, inhabiting today the the Pleistocene (for details see Szyndlar 2005, and refer- southeastern parts of the European continent. ences therein). Genus Telescopus Wagler, 1830. Type species: Telescopus obtusus Reuss, 1834. ?Telescopus sp. ‘‘Colubrinae’’ indet. Material: Two trunk vertebrae from Sandelzhausen. Material: Description: Ten trunk vertebrae and two maxillary fragments from The vertebrae display a number of peculiarities char- Sandelzhausen; seven trunk vertebrae from Rothenstein 13. acteristic of the living genus Telescopus, among others its Comments: parapophyses are distinctly longer than diapophyses, the These fragmentary vertebrae represented tiny snakes neural spine is extremely low, and the prezygapophyseal (with vertebral centra approximately 3 mm long), are processes are short and obtuse. Unfortunately, exact gen- morphologically different from the large-sized ‘‘Coluber’’ eric allocation based on such material (two poorly and ?Telescopus found in Sandelzhausen. preserved and fragmentary vertebrae only) is not possible. The vertebrae belonged to at least two different taxa. Comments: One vertebra from Sandelzhausen has a broad haemal keel The living genus Telescopus is widely distributed in in ventral view; the remaining vertebrae, coming from both Eurasia and Africa; one species, T. fallax, inhabits today sites, have very thin keels. All vertebrae are poorly pre- southeastern Europe. Fossil remains of Telescopus were served and devoid of protruding structures, so it is difﬁcult

123 60 Z. Szyndlar to ascertain whether the snakes from Sandelzhausen and of these snakes in the European continent began in the Rothenstein 13 represented different species or not. second half of the Early Miocene (MN3) (Ivanov 2002). Two small maxillary fragments from Sandelzhausen, Along with ‘‘colubrines’’, ‘‘natricines’’ (represented by at with partly preserved prefrontal processes, are only tem- least three different genera) were the most diverse and most porarily classified as belonging to ‘‘colubrines’’. widespread ophidian group in the period between the late Subfamily ‘‘Natricinae’’. Early and early Late Miocene (MN4 to 9). The genus Genus Natrix Laurenti, 1768. Natrix was represented in the aforementioned period by six Type species: Coluber natrix Linnaeus, 1758. extinct species. See Rage and Bailon (2005) and Szyndlar Natrix sp. (2005) for up-to-date reviews of the fossil record of Mio- Material: cene Natrix. Thirty-two trunk vertebrae from Sandelzhausen and ten cf. Natrix sp. (large form). trunk vertebrae from Rothenstein 13. Material: Description: Three trunk vertebrae from Sandelzhausen. All vertebrae are fragmentary and most of their pro- Comments: truding elements are lacking. For instance, only one Morphologically, these vertebrae do not differ consid- vertebra has fully preserved hypapophysis, whereas only erably from those referred above to Natrix sp., but they two vertebrae have preserved prezygapophyseal processes must have belonged to (a) specimen(s) of distinctly larger (on one side only). The vertebral centra are considerably dimensions, perhaps representing a distinct species. The elongated, with flat ventral surface delimited laterally with centrum length measured in two vertebrae is 7.4 mm. The usually strongly developed subcentral ridges. The centrum bones are badly damaged. length ranges between 3.0 and 4.0 mm. The hypapophysis Genus Neonatrix Holman, 1973. (preserved only in one vertebra from Sandelzhausen) is Type species: Neonatrix elongata Holman, 1973. rather straight (and not sigmoid-shaped) in lateral view and cf. ‘‘Neonatrix’’ sp. its distal tip is obtuse. The neural arch is weakly vaulted. Material: The neural spine (present partly in three vertebrae) is Four trunk vertebrae from Sandelzhausen and ten trunk somewhat lower than long, distinctly overhanging posteri- vertebrae from Rothenstein 13. orly (anterior portions of all spines are broken). The Description: paradiapophyses are strongly built, provided with promi- These vertebrae are relatively small, with the centrum nent parapophyseal processes, obtuse in shape and directed length 2.7 mm or shorter. The bones are poorly preserved; anteriorly; the diapophyses are usually as long as parap- however, most of them have retained complete (or almost ophyses. The zygosphenal roof is usually slightly convex in complete) neural spines and hypapophyses. The spines are dorsal view. The prezygapophyseal articular facets are oval- extremely low, whereas the hypapophyses are very short. shaped and elongate. Few postzygapophyseal articular Comments: facets are preserved; they are small and subsquare. The Both these characteristics were considered the most prezygapophyseal processes (missing in most vertebrae) are important diagnostic features of the extinct genus Neona- as long as the prezygapophyseal articular facets and slender. trix (cf. Rage and Holman 1984). Later, however, in the The cotyle and condyle are orbicular. The subcentral, lateral emended generic diagnosis of this genus, Auge´ and Rage and paracotylar foramina are distinct, though rather small. (2000) redefined Neonatrix as ‘‘a small »natricine« with a Comments: short hypapophysis’’ (whereas the significance of a low The morphological characteristics of the discussed ver- neural spine was ignored), an action criticised by Szyndlar tebrae are typical of the living genus Natrix. One of the (2005) (cf. also Szyndlar and Schleich 1993). most important diagnostic features is the shape of the hy- In the opinion of Szyndlar and Schleich (1993), this papophysis, in our case unfortunately preserved in one genus may be an artificial, polyphyletic taxon; for this vertebra only. Although the hypapophysis is not sigmoid- reason, the generic name Neonatrix is presented in quota- shaped (as characteristic of most vertebrae of members of tion marks. the genus Natrix), straight-shaped hypapophyses are also Snakes classified in the genus Neonatrix, originally not uncommon in trunk vertebrae of Natrix (especially in described from the Miocene of North America, were those located in more anterior portions of the column). widespread in the European continent in the period Identification of Natrix from Sandelzhausen and Rothen- between the late Early Miocene (MN4) (Szyndlar 1987; stein 13 to the specific level, owing to the poor Szyndlar and Schleich 1993; Rage and Bailon 2005) and preservation of the vertebrae, is not possible. the beginning of the Pliocene (MN14) (cf. Neonatrix; Members of the genus Natrix were present in Europe Venczel 2001). See Holman (2000) for the review of the since the Early Oligocene (Rage 1988), but the great bloom North American record of the genus. 123 Snake fauna from the Early/Middle Miocene of Sandelzhausen and Rothenstein 13 61

‘‘Natricinae’’ indet. indentation between the pre- and postzygapophyses. The Material: centrum length is 7.7 mm, centrum width is 6.5 mm; it is Fifty trunk vertebrae from Sandelzhausen. triangular in ventral view, with almost ﬂat ventral surface. Comments: The subcentral ridges are well developed. The hypapoph- These vertebrae cannot be identiﬁed owing to their ysis and neural spine remain unknown. The neural arch is strong damage. Probably they belonged to the aforemen- weakly vaulted. The paradiapophyses are well developed; tioned natricine snakes. the parapophyseal processes are projected anteriorly and Family Elapidae Boie, 1827. obtuse in shape. The zygosphenal roof is roughly straight in Genus Naja Laurenti, 1768. dorsal view, laterally provided with two indistinct lobes. Type species: Coluber naja Linnaeus, 1758. The prezygapophyseal articular facet (preserved in another Naja sp. mid-trunk vertebra: Fig. 3e) is oval-shaped. The prezyga- Figure 3. pophyseal process is well developed, somewhat shorter than Material: the articular facet a, having moderately obtuse tip; it is Six trunk vertebrae from Sandelzhausen and (possibly) directed laterally. The cotyle and condyle are orbicular. The eight trunk vertebrae from Rothenstein 13. subcentral, lateral and paracotylar foramina are distinct. Description: One vertebra coming from the posterior (but not poste- The vertebrae from Sandelzhausen come from the mid- riormost) trunk portion of the column (Fig. 3f–g) is much dle and posterior portions of the trunk region of the column. more elongate than the mid-trunk vertebrae. The hypap- The largest and best preserved middle trunk vertebra ophysis is very short, triangular-shaped, with the distal tip (Fig. 3a–d) is incomplete; its protruding parts are lacking. It acute and projecting posteriorly. The neural spine is very is relatively very short, with rather weakly developed low. It does not differ substantially from the posterior trunk

Fig. 3 Four trunk vertebrae (a–d BSPG 1959 II 16096; e BSPG 1959 II 16097; f, g BSPG 1959 II 16018); h, i BSPG 1959 II 15958) of Naja sp. from Sandelzhausen. a, f, h right lateral views; b ventral view; c, e, g, i dorsal views; d posterior view 123 62 Z. Szyndlar vertebrae of Naja romani from the French Middle Miocene Comments: depicted by Hoffstetter (1939, pl. II Fig. 8; originally The systematic allocation of these vertebrae is unclear. described as Palaeonaja romani). Most likely they belonged to elapid snakes, but their Another vertebra from Sandelzhausen (Fig. 3h–i) may allocation in the colubrid subfamily ‘‘Natricinae’’ is also have been the last trunk vertebra (or one of a few last trunk possible. vertebrae). It lacks the neural spine and left prezygapoph- The discussed vertebrae resemble those of a number of ysis, but besides it is perfectly preserved. The centrum small elapids reported from several Miocene localities of length is 6.1 mm and centrum width is 4.5 mm. Its most Europe. Of them, the only snake identified to the specific striking feature is the shape of the hypapophysis: in lateral level is Micrurus gallicus (Rage and Holman 1984), a view, it looks like a subsquare-shaped plate. fossil member of the extant North American genus reported The vertebrae from Rothenstein 13, belonging to (a) from several Early and Middle (MN4 to MN7 ? 8) Mio- smaller individual(s), are badly damaged and their cene sites (see Rage and Bailon 2005, and references assignment to the genus cannot be fully demonstrated. therein). However, the vertebrae from Sandelzhausen and Comments: Rothenstein 13 seem to be morphologically closer to the The vertebrae are similar to those of Naja romani, the remains of another tiny alleged elapid snake first reported elapid widespread in Europe from the Early (MN4) to the from the Early Miocene (MN4) of Petersbuch 2 by Late (MN11) Miocene (Szyndlar and Rage 1990), and Szyndlar and Schleich 1993 (termed ‘‘Elapidae indet.’’); perhaps they actually belong to this species. However, these authors compared the snake from Petersbuch 2 with considering the poor preservation of the material (in par- the Asiatic living genus Maticora (now synonymised with ticular the mid-trunk vertebrae), this cannot be stated with Calliophis). Similar small elapids are known from Europe certainty. The bone interpreted as the last trunk vertebra from the period ranging in age from the Early (MN3) (Fig. 3h–i) is strikingly similar to that of N. romani (Ivanov 2000) to Late (MN9) Miocene (Szyndlar 2005), reported recently (Szyndlar 2005) from the Hungarian but their taxonomic status and possible relationships with locality of Rudabańya. Szyndlar made attempts to recon- other fossil and living snakes remain unknown. struct the morphology of the posterior trunk portion of the McDowell (1987) once suggested close similarities vertebral column of N. romani, pointing out similarities between New World (Micrurus) and Asian Coral snakes between this fossil species and its living relative, N. naja (Calliophis). However, Slowinski et al. (2001) demon- from southern Asia. It is not certain, however, whether the strated that Asian coral snakes (Calliophis) are middle trunk and posterior trunk vertebrae from Sandelz- paraphyletic relatives to New World coral snakes (Micru- hausen belonged to the same species of cobra. The snake rus). They proposed that the origin of New World coral from which the posteriormost vertebra originates must snakes was Asian; hence, New World coral snakes derived have possessed much larger mid-trunk vertebrae than those from an ancestor which arrived into the Americas via found in Sandelzhausen. Also the cobra from Rothen- Bering Land Bridge. stein 13 may have represented another species, Family Viperidae Oppel, 1811. characterized by smaller dimensions. Genus Vipera Laurenti, 1768. For the full account of the fossil record of the genus Type species: Coluber aspis Linnaeus, 1758. Naja in Europe see Szyndlar and Rage (1990); for recent Vipera sp. (‘‘aspis complex’’). updates see Szyndlar (2005). Material: Elapidae indet. Twenty trunk vertebrae and one right compound bone Material: from Rothenstein 13. Six trunk vertebrae from Sandelzhausen and two trunk Description and comments: vertebrae from Rothenstein 13. Members of the living genus Vipera referred to the so- Description: called aspis complex were present in Europe, including These fragmentary vertebrae, belonging to relatively Germany, since the beginning of the Miocene (MN1) small snakes, are characterized by elongated centra and (Szyndlar and Bo¨hme 1993). Since the end of the Early very short (vestigial) hypapophyses. The latter structures, Miocene (MN4), in most fossil assemblages these reptiles completely preserved in most vertebrae, are the only pro- were usually accompanied by members of another complex truded elements in the discussed remains. The centrum of the genus Vipera, namely ‘‘Oriental vipers’’ (Szyndlar length of most vertebrae found in both localities ranges and Rage 2002). Specific identification of viperid fossils is between 3.0 and 4.0 mm. In one vertebra from Sandelz- very difficult, but distinguishing the two complexes based hausen (belonging to an adult snake) the centrum length is on vertebrae is a relatively easy task. Vertebrae of the 1.5 mm only. It is possible that the vertebrae belonged to ‘‘Oriental vipers’’ are relatively very large and short, hav- two or more different species. ing high neural spines, whereas those of the ‘‘aspis 123 Snake fauna from the Early/Middle Miocene of Sandelzhausen and Rothenstein 13 63 complex’’ are smaller and more elongate, having lower Description and comments: neural spines. The presence of the ‘‘Oriental vipers’’ in Sandelzhausen The viperid vertebrae from Rothenstein 13, although was first reported by Szyndlar and Schleich (1993). Most strongly damaged, display clearly the conditions charac- vertebrae, coming from the middle trunk portion of the teristic of the latter group. The centrum length of the column, although badly damaged, display clearly charac- largest vertebral fragment is 6.0 mm; the remaining ver- teristic of the ‘‘Oriental vipers’’. The centrum length of the tebrae have smaller dimensions. largest vertebra is 9.2 mm; centrum width is 8.5 mm; A small posterior fragment of a compound bone (with- centrum length/centrum width ratio \1.1. out number in the collection of the SMNS) is identifiable as The centrum of the trunk vertebrae is distinctly trian- belonging to Vipera based on its relatively high medial gular-shaped in ventral view. The subcentral ridges and flange. subcentral grooves are distinct in the anterior portion of Fossil remains of Vipera belonging to the ‘‘aspis com- the centrum. The hypapophyses are missing in all verte- plex’’ were fairly common in Miocene and Pliocene sites. brae, but the remnants indicate that the structure was very Today the range of the distribution of these vipers is strong and thick. The neural arch is strongly depressed restricted almost exclusively to southern Europe (south of and tilted in the postzygapophyseal area, devoid of epi- the Alps). See Szyndlar and Rage (2002) for more infor- zygapophyseal spines. The neural spine is lacking in all mation on the past distribution of ‘‘aspis-like’’ vipers and vertebrae. The zygosphene is very thick in anterior view other members of the genus Vipera (s.l.). and straight in dorsal view. The prezygapophyseal artic- Vipera sp. (‘‘Oriental viper’’). ular facet, preserved on one side in one vertebra only, is Figure 4. rectangular-shaped, strongly elongated laterally. Prezyga- Material: pophyseal processes, parapophyses and parapophyseal Eighteen trunk vertebrae, one cloacal vertebra and [50 processes are either lacking or eroded in all available isolated venomous fangs from Sandelzhausen. vertebrae. The cotyle and condyle are (usually strongly)

Fig. 4 Four trunk vertebrae (a BSPG 1952 II 16090; b BSPG 1952 II viper’’) from Sandelzhausen. a right lateral view; b, d, i anterior 16091; c–e BSPG 1959 II 16010; f–h BSPG 1959 II 16095) and a views; c left lateral view; e, g posterior views; f dorsal view; h ventral cloacal vertebra (i BSPG 1959 II 15976) of Vipera sp. (‘‘Oriental views 123 64 Z. Szyndlar

flattened dorso-ventrally. All vertebral foramina are the different faunal composition might be due to tapho- distinct. nomical effects, where a higher specimen number supports Today ‘‘Oriental vipers’’ (in recent herpetological liter- the probability of a record of a higher taxonomical diver- ature usually classified in the genera Macrovipera and sity. However, the fossil remains are too scarce and too Montivipera) are distributed in Asia and northern Africa; in fragmentary to give answers on these questions. But, since Europe, their distribution is restricted to southeastern out- differences in the habitat conditions are indisputable, pos- skirts of the continent. However, in the Neogene they sible differences in the adaptations of single taxa have to be inhabited vast areas of Europe (at least its southern half) considered, especially in creeping reptiles, for which this and belonged to the commonest elements of fossil snake study might be an initial suggestion. assemblages. See Szyndlar and Rage (2002) for more The most interesting component of the discussed snake information on the past distribution of these vipers. faunas is the extinct aniliid genus Eoanilius: Sandelzhau- Serpentes indet. sen is the geologically youngest site yielding this Eocene Material: survivor. Similarly, the extinct boid Bavarioboa from Ro- One hundred and thirty caudal vertebrae, 2 dentary thenstein 13 (along with a related snake from the coeval fragments and 1 fragment of an unidentified toothed bone locality of Isle d’Abeau in France) is the geologically from Sandelzhausen; 46 trunk vertebrae, 13 caudal verte- youngest representative of the genus (Szyndlar and Rage brae and 2 ?pterygoid fragments from Rothenstein 13. 2003). Comments: Fossil snakes do not seem to be a useful tool for The bones likely belonged to the above described taxa, reconstructing most factors of palaeoecosystems; never- but this cannot be demonstrated owing to the poor pres- theless, they are commonly considered very good ervation of the remains. A small fragment of an indicators of past climates, in particular environmental unidentified toothed bone (bearing two teeth) and two temperatures. This property results directly from their fragmentary dentaries may have belonged to colubrids. physiological dependence on external sources of heat. Although the reduction in diversity from lower to higher latitudes can be observed in virtually all groups of animals Discussion and plants worldwide, among terrestrial vertebrates reptiles (including snakes) are particularly sensitive to cold. The overall composition of the assemblages from San- In other words, their south-to-north geographical distri- delzhausen and Rothenstein 13 is roughly the same as bution is largely determined by available heat energy. The those reported from two late Early Miocene (MN4) local- composition of the snake fauna from Sandelzhausen and ities situated nearby: Dolnice in western Czechia (Szyndlar Rothenstein 13, similarly to the older (MN4) faunas 1987) and Petersbuch 2 in southern Germany (Szyndlar coming from several nearby localities, indicates that air and Schleich 1993). Unfortunately, owing to the poor temperatures in the area must have been distinctly higher preservation of the fossils from Sandelzhausen and Ro- that today. However, the climate in Central Europe must thenstein 13, most faunal elements cannot be compared have been less favourable for highly thermophilous rep- precisely at the specific level. At the generic level, the tiles than in the case of more southern parts of Europe. fossils do not differ substantially from those found in a The most obvious indicator of lower temperatures in number of European sites representing European Land southern Germany compared with southern Europe in the Mammal Zones MN4 and MN6 (cf. Ivanov 2000, fig. 16; Miocene is the absence of the genus Python successfully Rage and Bailon 2005, tab. 1). invading Mediterranean coasts at the Early/Middle Mio- The snake assemblage from Sandelzhausen differs from cene transition (Szyndlar and Rage 2003). Other obvious that of Rothenstein 13 by the presence of Eoanilius, candidates for highly thermophilous forms could be Telescopus and ‘‘Oriental viper’’. In contrast, the assem- cobras of the genus Naja. Unfortunately, our knowledge blage from Sandelzhausen lacks Bavarioboa and the about possible relationships between the extinct cobras ‘‘aspis-like’’ Vipera, which are represented in Rothen- from Sandelzhausen and Rothenstein 13 and their living stein 13. These differences in the faunal compositions relatives from Asia is limited, again due to poor preser- might indicate the ecologically different habitats (wet vation of the fossil remains. Most living members of the Molasse Basin and dry karstic area of the Franconian Alb) genus Naja inhabiting the Asiatic continent today are (Kaiser and Ro¨ssner 2007) or ecologically differently highly thermophilous; N. oxiana, however, distributed adapted species. The main ecological difference of both relatively closely to the European eastern outskirts and habitats are seasonally dry conditions in the Molasse Basin reaching the elevations of almost 2,000 m (Joger 1984), is compared with year-round dry edaphic conditions and apparently an animal adapted to more severe climatic missing open water in the karstic area. On the other hand, conditions. 123 Snake fauna from the Early/Middle Miocene of Sandelzhausen and Rothenstein 13 65

Conclusion Kaiser, T., and G.E. Ro¨ssner. 2007. Dietary resource partitioning in ruminant communities of Miocene wetland and karst palaeoenvironments in Southern Germany. Palaeogeography, The ophidian faunas from Sandelzhausen and Rothen- Palaeoclimatology, Palaeoecology 252(3–4):424–439. stein 13 fill a gap in knowledge between MN4 and MN6, McDowell, S.B. 1987. Systematics. In Snakes: Ecology and Evolu- which were expected to document a turnover from still tionary Biology, eds. Seigel, R.A., Collins, J.T., and Novak, S.S., archaic influenced to completely modern composed com- 3–50. New York: Macmillan. Milner, A.C., A.R. Milner, and R. Estes. 1982. Amphibians and munities. In fact, the hitherto youngest known occurrences squamates from the Upper Eocene of Hordle Cliff, Hampshire— of the last archaic elements Eoanilius and Bavarioboa a preliminary report. Tertiary Research 4(4):149–154. could be identified, demonstrating a successive moderni- Moser, M., Ro¨ssner, G.E., Go¨hlich, U.B., Bo¨hme, M., and Fahlbusch, sation of Central European ophidian faunas from the late V. 2009. this volume. The fossil lagersta¨tte Sandelzhausen (Miocene, southern Germany): history of investigation, geology, Early to the early Middle Miocene. fauna and age. In Fossil lagersta¨tte Sandelzhausen (Miocene, Based on the ophidian fossils, the climates of Sandelz- southern Germany): Contributions to the fauna. Palaöntologi- hausen and Rothenstein 13, although warm, cannot be sche Zeitschrift, eds. Ro¨ssner, G.E., and Go¨hlich, U.B., 83 (1), considered as tropical or subtropical. 000–000. Rage, J.C. 1974. Les serpents des Phosphorites du Quercy. Palae- overtebrata 6(3–4):274–303. Acknowledgments I am deeply grateful to Prof. Volker Fahlbusch Rage, J.C. 1988. The oldest known colubrid snakes. The state of the and Dr. Gertrud Ro¨ssner for inviting me to participate in the San- art. Acta Zoologica Cracoviensia 31(13):457–474. delzhausen project as well as for all courtesies received from them in Rage, J.C., and S. Bailon. 2005. Amphibians and squamate reptiles Mainburg during the ‘‘Sandelzhausen Symposium 2005’’. I thank Dr. from the late Early Miocene (MN4) of Beón 1 (Montreál-du- Ronald Bo¨ttcher (Staatliches Museum fu¨r Naturkunde in Stuttgart) for Gers, southwestern France). Geodiversitas 27(3):413–441. the loan of the material from Rothenstein 13. Dr. Massimo Delfino Rage, J.C., and J.A. Holman. 1984. Des Serpents (Reptilia, Squamata) (Florence) and Dr. Jean-Claude Rage (Paris) provided helpful com- de type Nord-Ame´ricain dans le Mioce`ne Français. E´ volution ments on the manuscript. Dr. Dave Hone made linguistic corrections paralle`le ou dispersion? Geóbios 17(1):89–104. of the text. Rage, J.C., and Z. Szyndlar. 2005. 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