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"Dinosauromorph" Tracks from the Middle Buntsandstein (Early Triassic: Olenekian) of Wolfhagen, Northern Hesse, Germany

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Versão online: http://www.lneg.pt/iedt/unidades/16/paginas/26/30/141 Comunicações Geológicas (2013) 100, 1, 81-88 ISSN: 0873-948X; e-ISSN: 1647-581X "Dinosauromorph" tracks from the Middle Buntsandstein (Early Triassic: Olenekian) of Wolfhagen, northern Hesse, Germany Pistas "Dinossauromórficas" do Médio Buntsandstein (Triásico Basal: Olenekiano) de Wolfhagen, Hesse norte, Alemanha J. Fichter1,*, R. Kunz2 Recebido em 30/04/2012 / Aceite em 26/07/2012 Artigo original Disponível online em Julho de 2012 / Publicado em Junho de 2013 Original article © 2013 LNEG – Laboratório Nacional de Geologia e Energia IP Abstract: Exposures of the Detfurth Formation of the Middle preparation. The present study is concerned with footprints referable Buntsandstein (Early Triassic: Olenekian) in an abandoned sandstone to the ichnogenera Prorotodactylus and Rotodactylus. quarry near Wolfhagen (northern Hesse, Germany) have yielded Brusatte et al. (2010) reported small reptilian tracks from the numerous reptilian tracks referable to the ichnogenera Prorotodactylus Olenekian of the Swietokrzyskie (Holy Cross) Mountains (Poland) and Rotodactylus. These specimens are described here and compared to footprints of the ichnogenus Rhynchosauroides, which are common in the and referred them to the ichnogenus Prorotodactylus, which same stratigraphic interval. superficially resembles Rhynchosauroides. Based on comparative analysis they argued that the Polish tracks were produced by Keywords: Prorotodactylus, Rotodactylus, tetrapod footprints, Triassic, Germany. dinosauromorph archosaurs that were closely related to Lagerpeton, which is known from skeletal remains from the Middle Triassic Resumo: Exposições da Formação Detfurth do Buntsandstein Médio (Ladinian) Chañares Formation of northwestern Argentina (Romer, (Triásico Basal: Olenekiano) numa pedreira de arenitos abandonada perto 1971; Sereno and Arcucci, 1994). Thus, Brusatte et al. (2010) de Wolfhagen (Hesse Norte, Alemanha) forneceram numerosas pistas interpreted the Polish finds as evidence for extending the known reptilianas referentes aos icnogéneros Prorotodactylus e Rotodactylus. stratigraphic range of Dinosauromorpha back to the Early Triassic. Estes espécimes são descritos e comparados com as pegadas do This surprising discovery led the authors to examine in detail icnogénero Rhynchosauroides, que são comuns no mesmo intervalo track-bearing slabs from the Detfurth Formation near Wolfhagen to estratigráfico. search for similar tracks. During the first half of 2011, additional, Palavras-chave: Prorotodactylus, Rotodactylus, pegadas tetrápodes, very well preserved footprints referable to Prorotodactylus and Triásico, Alemanha. Rotodactylus from the talus slope of the quarry face were collected. 1Jürgen Fichter Martinsplatz 4, D-34117 Kassel, Germany The new material was examined and documented, and in subsequent 2Reiner Kunz Friedensstrasse 7, D-34466 Wolfhagen, Germany discussion with H.-D. Sues, it turned out that there were unresolved *Corresponding author/Autor correspondente: [email protected] issues in the zoological interpretation of the track makers. 1. Introduction 2. Geographic and stratigraphic context In 1999 R. Kunz and J. Fichter discovered tracks of reptiles in an The locality (Fig. 1) for the material described in this paper is a abandoned sandstone quarry in the municipal forest at Wolfhagen sandstone quarry that has not been actively worked for about 15 (northern Hesse), which exposes sections of the Detfurth Formation years. It is located about 2 km southwest of the small town of of the Middle Buntsandstein (Early Triassic: Olenekian) (Fichter et Wolfhagen in northern Hesse, almost in the geographic center of al., 1999; Kunz and Fichter, 2000; Fichter and Kunz, 2006). Since conterminous Germany. that time numerous track-bearing slabs have been collected both The stratigraphic sequence (Fig. 2) exposed in the quarry during excavations in the exposed strata and during prospecting in comprises the top of the Detfurth Sandstone and the lower and the talus slope at the bottom of the quarry face. To date nine middle parts of the approximately 20 to 25 m thick Detfurth ichnotaxa have been recorded: ?Capitosauroides, Dicynodontipus, "Wechselfolge" of the Middle Buntsandstein (Horn, 1976; Fichter et Procolophonichnium, Prorotodactylus, Protochirotherium, al., 1999). The age of these strata is Olenekian (see Brusatte et al., Palaeochirotherium, Rhynchosauroides, Rotodactylus, and 2010). The sequence is characterized by a succession of colored Synaptichnium. claystones and fine- to medium-grained, predominantly reddish Fichter and Kunz (2004) first introduced the chirotherioid brown sandstones. Almost all of the mudstone beds preserve ichnogenus Protochirotherium, which is now widely used (Klein and mudcracks, which indicate repeated drying out of the depositional Haubold, 2007; Klein et al., 2010; Klein and Lucas, 2010; Tourani et setting. All of the footprints described in this paper were recovered al., 2010; Avanzini et al., 2011). More recently, they named a second from this sequence, of which about 12 m are exposed in the quarry. new ichnogenus, Palaeochirotherium, based on tracks found in 2010 They are preserved on the underside of sandstone beds, which (Fichter and Kunz, 2011). A comprehensive survey of the tetrapod directly overlie mudstones. Fichter et al. (1999) provided additional ichnofauna from the Detfurth Formation of Wolfhagen is currently in details concerning the lithology and depositional setting. 82 J. Fichter et al. / Comunicações Geológicas (2013) 100, 1, 81-88 Fig.1. General geographic position of Wolfhagen. Fig.1. Localização geográfica de Wolfhagen. Fig.2. Profile and stratigraphy of the Wolfhagen quarry tracksite as well as representation of the ichnogenera found so far. Fig.2. Perfil e estratigrafi da pedreira Wolfhagen, bem como uma representação dos icnogéneros encontrados até agora. 3. Systematic Palaeontology from the base to the tip of pedal digit IV because the more proximal portion of the foot has not been impressed. The tracks generally Ichnofamily Prorotodactylidae Ptaszyiński, 2000 preserve only impressions of pedal digits II-IV. Occasionally traces Ichnogenus Prorotodactylus Ptaszyiński, 2000. of the first and fifth digit are visible. The angle between digits II and Material: RGMW-SK-WSF-1-0, RGMW-SK-WSF-30-1, IV is usually less than 10° so that the three central digits extend RGMW-SK-WSF-38-0, RGMW-SK-WSF-38-1, RGMW-SK-WSF- nearly parallel to one another. If at all preserved the proximal edge of 39-2, RGMW-SK-WSF-53-0, RGMW-SK-WSF-53-1, RGMW-SK- the pedal imprint is straight. The toes distinctly increase in length WSF-53-2, RGMW-SK-WSF-53-3, RGMW-SK-WSF-59, RGMW- from the first to the fourth. In well-preserved imprints the toes clearly SK-WSF-60-2, RGMW-SK-WSF-60-3, RGMW-SK-WSF-60-4, show distinct, narrow plantar scales, which are aligned at right angles RGMW-SK-WSF-63, RGMW-SK-WSF-64, RGMW-SK-WSF-66- to the long axes of the toes (Fig. 3 A, B; Fig. 4 E; see also 0, RGMW-SK-WSF-66-1, RGMW-SK-WSF-67, RGMW-SK- Ptaszyiński, 2000). Sometimes overprinting of tracks can be WSF-68-0, RGMW-SK-WSF-71-0, RGMW-SK-WSF-76 (not all observed. illustrated here; see Fig. 4). Manus impressions: On several specimens (e.g., Fig. 4 I, J, K) Labelling of specimens is similar to the method of Fuglewicz et manus impressions probably associated with pes impressions have al. (1990). It consists of an acronyme of the collection and a number been more or less clearly preserved. They appear more compact and of the slab and of the footprint on this slab. For example RGMW- distinctly smaller than the pes impressions. Ptaszyiński (2000) noted SK-WSF-66-2 means, that the footprint nº 2 on the slab nº 66 is that the third rather than the fourth manual digit is the longest in stored at the “Regionalmuseum Wolfhager Land” under “Sammlung- Prorotodactylus. Specimen RGMW-SK-WSF-76 (Fig. 4 I) appears to Kunz-Wolfhager-Saurierfährten”. confirm this interpretation. However, one must consider the possibility Pes impressions: The pes impressions are digitigrade, five-toed, that the distal portion of manual digit IV was overprinted by the pes and range in length from about 20 to slightly more than 30 mm. It impression. The angle between manual digits II and IV (about 16° to should be noted that these measurements only represent the distance 19°) is slightly larger than that on the pedal impressions. "Dinosauromorph" tracks of Wolfhagen, Germany 83 Fig.3. Pes imprint of Prorotodactylus isp. showing distinct, narrow plantar scales, which are aligned at right angles to the long axes of the toes (SK-WSF-66-2). A detail photo, B graphical interpretation. Scale = 1 cm. Fig.3. Impressão Pes do Prorotodactylus isp. mostrando distintas e estreitas escamas plantares que estão alinhadas a ângulos direitos com os eixos longos dos dedos (SK- WSF-66-2). A foto detalhada, B interpretação gráfica. Escala = 1 cm. Fig.4. Prorotodactylus isp., photos and corresponding drawings: A-H pes impressions without associated manus impressions [A (SK-WSF 39-2), B (SK-WSF-1- 0), C (SK-WSF 63), D (SK-WSF-66-1), E (SK-WSF- 66-2), F (SK-WSF-60-2), G (SK-WSF-60-3), H (SK- WSF-64)]. I-K pes impressions (p) probably associated with manus impressions (m) [I (SK-WSF-76), J (SK- WSF-68-0), K (SK-WSF-67)]. L pes impression (SK- WSF-59). M isolated, exceptionally large ?manus impression (SK-WSF-71-0). N isolated, very small ?manus impression (SK-WSF-30-1). O manus impression (detail of track I). P reconstructed complete foot morphology of Prorotodactylus. Scale 1 = cm. Fig.4. Fotos de Prorotodactylus isp. e desenhos correspondentes: A-H impressões pes com as impressões manus associadas [A (SK-WSF 39-2), B (SK-WSF-1-0), C (SK-WSF 63), D (SK-WSF-66-1), E (SK-WSF-66-2), F (SK-WSF-60-2), G (SK-WSF-60-3), H (SK-WSF-64)]. I-K impressões pes (p) provavelmente associadas com impressões manus (m) [I (SK- WSF-76), J (SK-WSF-68-0), K (SK-WSF-67)]. L impressões pes (SK-WSF-59).
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