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.
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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). M impressões manus isoladas, excepcionalmente largas? (SK-WSF-71-0). N impressões manus isoladas muito pequenas? (SK-WSF-30-1). O impressões manus (detalhe da pista I). P morfologia completa reconstruída do pé de Prorotodactylus. Escala = 1 cm.
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Ichnofamily Rotodactylidae Peabody, 1948 the fourth digit. The angles between digits II, III, and IV are Ichnogenus Rotodactylus Peabody, 1948 comparable to those in Prorotodactylus. The first pedal digit is only faintly visible in exceptional circumstances (Fig. 5 F). The Material: RGMW-SK-WSF-39-0, RGMW-SK-WSF-41-0, fifth digit is represented by a slight impression of its tip behind and RGMW-SK-WSF-48-0, RGMW-SK-WSF-60-0, RGMW-SK- slightly lateral to the impression of the fourth digit. WSF-60-1, RGMW-SK-WSF-66-3, RGMW-SK-WSF-70, RGMW-SK-WSF-71-0, RGMW-SK-WSF-71-1, RGMW-SK- Manus impressions: We interpret a few impressions referred WSF-71-2 (not all illustrated; see Fig 5). to Rotodactylus as having been produced by the manus (e.g. Fig. 5 Pes impressions: The pes impressions are five-toed and I). They are distinctly smaller but closely resemble the pes distinctly digitigrade, with average lengths ranging from 30 to 50 impressions except for the occasionally more pronounced mm. Pedal digits II, III, and IV are uniformly impressed and, as in divergence between the first four digits and the more lateral Prorotodactylus, show a distinct increase in length from the first to position of the imprint of the tip of the fifth digit.
Fig.5. Rotodactylus isp., photos and corresponding drawings: A-H pes impressions, [ A (SK-WSF-39-0), B (SK-WSF-41-0), C (SK-WSF-60+1), D (SK-WSF-60-1), E (SK-WSF 60- 0), F (SK-WSF-66-3), G (SK-WSF 70), H (SK WSF-71-0 )]. I manus impression (SK-WSF-71-1]. Scale 1 = cm.
Fig.5. Fotos de Rotodactylus isp. e desenhos correspondentes: A-H impressões pes, [A (SK-WSF-39-0), B (SK-WSF-41-0), C (SK-WSF-60+1), D (SK-WSF-60-1), E (SK-WSF 60- 0), F (SK-WSF-66-3), G (SK-WSF 70), H (SK WSF-71-0)]. I impressões manus (SK-WSF-71-1]. Escala = 1 cm..
4. Comparison with Rhynchosauroides Prorotodactylus and Rotodactylus differ from Rhynchosauroides in the distinctly parallel orientation of pedal The footprints of Prorotodactylus and Rotodactylus closely digits II-IV. This feature is even more pronounced in resemble those of Rhynchosauroides (Fig. 6 A, B, C) and thus Prorotodactylus than in Rotodactylus. In both Prorotodactylus and further comparison and discussion are warranted. The three Rotodactylus the proximal delineation of digit impressions is ichnogenera represent what Nopcsa (1923) termed the lacertoid straight where preserved whereas this is not the case in type and which is characterized by a distinct increase in length Rhynchosauroides. Only in well-preserved tracks of Rotodactylus from the first to the fourth digit in both manus and pes impressions is there a clear trace of the tip of digit V behind digits I-IV. Tracks and the posterior position of the fifth digit. Thus, a clear distinction without this trace cannot be assigned to Rotodactylus. This unusual between the three ichnogenera is only possible for well and more retroversion of the fifth digit is absent in Prorotodactylus and or less completely preserved track. The completeness of especially Rhynchosauroides. In Prorotodactylus a posteriorly preservation depends on the position of the foot relative to the situated trace of the fifth digit is occasionally present lateral to digit substrate as well as other locomotion- and substrate-related factors. IV (Fig. 4 C, F, G). This indicates that pedal digit V extended Size differences are not critical because of the already-known straight forward. On a few exceptional, deeply imprinted tracks of ranges in size. Prorotodactylus parts of the sole and heel can be visible (Fig. 4 A, "Dinosauromorph" tracks of Wolfhagen, Germany 85
F, L). They suggest a foot with a rather narrow, posteriorly more or noteworthy (Fig. 8) whereas Prorotodactylus is only a minor element less conically tapering plantar surface (Fig. 4 M). in the ichnofaunas from the Stryczowice and Wióry sequences Based on the studies by Ptaszyiński (2000) and Brusatte et al. (Brusatte et al., 2010). Prorotodactylus has not yet been reported (2010) the manus impressions of Prorotodactylus clearly differ from from correlative strata in Morocco (Klein et al., 2010) and Italy the pes impressions in that manual digit III is the longest, the (Avanzini et al., 2011). divergence between digits is generally greater, and digit V is distinct Tracks of Rotodactylus are uncommon in the Wolfhagen from digits I-IV. This resembles Nopcsa's (1923) chirotherioid type ichnofauna. This ichnogenus is usually found only in rather than the lacertoid type represented by the pedal impressions stratigraphically slightly younger units. In Poland Rotodactylus is or tracks of Rhynchosauroides. Ptaszyiński (2000) discussed this known from the Baranow sequence, which corresponds to the Röt marked difference between manus and pes impressions and sequence of the Buntsandstein (early Anisian). Rotodactylus is also attempted to explain it as reflecting distinct specializations of the present at the transition from the Solling to the Röt sequence of the forelimb. Some of the pedal impressions referred to Prorotodactylus Buntsandstein in Thuringia (Haubold, 1967, 1971a-b, 1999), the and figured by Brusatte et al. (2010: Fig. 2 a, h, i) also resemble Solling sequence of Bad Karlshafen in northern Hesse (Demathieu those of the chirotherioid type. This raises the possibility that such and Fichter, 1989), the Middle Triassic of France (Demathieu, 1984), footprints simply represent small examples of Synaptichnium. One Italy (Avanzini et al., 2011; Todesco and Bernardi, 2011), Spain of the tracks from the Wolfhagen locality closely resembles the (Fortuny et al., 2011), Algeria (Kotanski et al., 2004), and Morocco reconstructed pes impression of Prorotodactylus (Fig. 4 M, P), but (Klein et al., 2011), and from the Moenkopi Formation of Arizona the impression of digit III appears to be slightly longer than that of (Peabody, 1948). Rhynchosauroides is the dominant ichnotaxon in digit IV and this specimen might represent an unusually large manus the footprint assemblage from the Detfurth sequence at Wolfhagen impression of Prorotodactylus. based on the total amount of material collected from the talus slope of the quarry face, which apparently represents several track horizons (Fig. 7). This also appears to be the case for other localities and a broader range of stratigraphic sequences. For tetrapod ichnofaunas from the Middle Triassic of northeastern Spain, Fortuny et al. (2011) noted that footprints of Rhynchosauroides outnumber those of Chirotherium, Isochirotherium, Procolophonichnium, Rotodactylus, and Synaptichnium (see also Todesco et al., 2008).
Fig.6. Comparison of Prorotodactylus (A) and Rotodactylus (B) pes impressions with Rhynchosauroides (C) pes-manus impression couple. Scale 1 = cm. Fig.7. (A) slab SK-WSF 39 with Rhynchosauroides, Prorotodactylus, Rotodactylus and chirotherioid footprints. (B) slab SK-WSF-66 with Prorotodactylus and Fig.6. Comparação das impressões pes de Prorotodactylus (A) e Rotodactylus (B) com Rotodactylus footprints. (C) plate SK-WSF 79 with mainly Rhynchosauroides o par de impressões pes-manus Rhynchosauroides (C). Escala = 1 cm. footprints. Scale 5 cm each.
Fig.7. (A) Laje SK-WSF 39 com pégadas Rhynchosauroides, Prorotodactylus, 4. Comparison with other Occurrences Rotodactylus e chirotherioide. (B) Laje SK-WSF-66 com pégadas Prorotodactylus e Rotodactylus. (C) Laje SK-WSF 79 sobretudo com pégadas Rhynchosauroides. The Wolfhagen tetrapod ichnofauna most closely resembles that Escala = 5 cm. from Wióry in the Swietokrzyskie (Holy Cross) Mountains of Poland. Both assemblages comprise the characteristic ichnogenera Protochirotherium (frequently identified as Brachychirotherium in Poland), Palaeochirotherium (in part identified as Isochirotherium in Poland), Synaptichnium, Rhynchosauroides, ?Capitosauroides, Procolophonichnium, therapsid tracks, and Prorotodactylus (Klein and Haubold, 2007; Klein and Lucas, 2010; Fichter and Kunz, 2011). The Wióry sequence correlates more or less with the Detfurth and Hardegsen formations based on paleomagnetic and biostratigraphic data (Brusatte et al., 2010). The Stryczowice sequence is stratigraphically slightly older, corresponding to the Volpriehausen sequence of the Middle Buntsandstein based on conchostracans and Fig.8. Numerical distribution of the ichnotaxa proved so far in the Detfurth-formation palynomorphs (Brusatte et al., 2010). All the units discussed here are of Wolfhagen (without ?Capitosauroides). Olenekian in age. Note that Prorotodactylus occurs fundamentally more frequent than Rotodactylus within the "Dinosauromorpha". The most frequent track element is Rhynchosauroides. To date Prorotodactylus has only been recorded from the Stryczowice and Wióry sequences of Poland and from the Middle Fig.8. Distribuição numérica dos icnotaxa encontrados até agora na Formação Detfurth Triassic of France (Demathieu, 1984; Ptaszyiński, 2000). The relative de Wolfhagen (sem ?Capitosauroides). De notar que Prorotodactylus ocorre mais abundance of this ichnogenus in the Wolfhagen assemblage is frequentemente que Rotodactylus dentro dos "Dinosauromorpha". O elemento de pista mais frequente é Rhynchosauroides. 86 J. Fichter et al. / Comunicações Geológicas (2013) 100, 1, 81-88
5. The Identity of the Prorotodactylus Track Maker impressions for both Prorotodactylus and Rotodactylus, and this might suggest a trend toward bipedalism. Haubold (1967, 1971a) As previously noted, Brusatte et al. (2010) interpreted the producer inferred predominantly quadrupedal locomotion for the producer of of Prorotodactylus tracks as a dinosauromorph archosaur. This the Rotodactylus tracks even though the body proportions of his assessment is based on synapomorphies, which Ptaszyiński (2000) hypothetical track maker suggest at least the possibility of did not discuss when he first distinguished Prorotodactylus. One bipedalism. synapomorphy hypothesized for archosaurian tracks is very narrow trackways with a pace angulation of about 130°. As the Wolfhagen material does not include complete trackways we cannot determine the presence of this feature. Other features postulated as synapomorphies for Dinosauromorpha by Brusatte et al. (2010), such as pronounced digitigrady, closely apposed and parallel- extending digits II-IV, reduction of digits I and V, the absence of metatarsal impressions (with exception of occasional deeply impressed tracks in softer substrate) as well as the typically straight posterior edge of the footprints, can be clearly recognized in the Wolfhagen footprints. Brusatte et al. (2010) explicitly compared the producer of Prorotodactylus to the small dinosauromorph Lagerpeton. However, there are morphological reasons for challenging this interpretation. The pes of Lagerpeton could not possibly produce a footprint with impressions of digits I and V (the latter of which comprises only a splint-like metatarsal) (Fig. 9 A, B); however, these impressions are clearly evident in Prorotodactylus. Indeed, it is doubtful that even pedal digit II would have left an impression. Fig.10. (A) reconstructed pedal skeleton for Prorotodactylus (Scale = 1cm), (B) pedal Furthermore, the proximal delineation of the pes cannot be straight skeleton of the protorosaurian archosauromorph Pamelaria (without scale). The red line represents the joint level between metatarsus and phalanges for the toes II-III. due to the arrangement of the metatarsals in Lagerpeton if the axis of flexion extended between proximal phalanges 1 and 2 (Fig. 9). Fig.10. (A) Esqueleto do pé reconstruído para Prorotodactylus (Escala = 1cm), (B) The footprints of Prorotodactylus indicate a pes in which digits I Esqueleto do pé do protorossauro archosauromorfo Pamelaria (sem escala). As and V are reduced but still sufficiently well developed to leave linhas vermelhas representam o nível da ligação entre o metatarso e as falanges para os dedos II-III. impressions. As the metatarsal region is not evident in the impressions the axis of flexion must have extended at least between the metatarsals and first phalanges of the digits II-IV. Figure 10 A Haubold (1998) noted that Rotodactylus is the most common illustrates a hypothetical pedal structure that corresponds to the ichnogenus in track-bearing horizons of the Lower Triassic. He tracks of Prorotodactylus. We note that this reconstruction also cited an example from the Solling Formation in Germany where resembles the pes of the protorosaurian archosauromorph up to 1,000 footprints of Rotodactylus per square meter have Pamelaria (Fig. 10 B) been recorded. In the stratigraphically older Detfurth Formation near Wolfhagen, however, Rotodactylus is uncommon, indeed much less common than Prorotodactylus, despite excellent preservational conditions. Klein and Haubold (2007) and Klein and Lucas (2010) proposed a biochronology of the continental Triassic based on tetrapod tracks. Klein and Haubold (2007) defined six biochrons on the basis of the first appearance datum (FAD) for a characteristic index ichnotaxon for each assemblage. More recently Klein and Lucas (2010) only recognized five zones. Biochron I is defined in its upper portion by the appearance of Protochirotherium (Fichter and Kunz, 2004) and corresponds to the late Induan to Olenekian. However, Avanzini et al. (2011) Fig.9. (A) pes skeleton of Lagerpeton chanarensis, (B) digitigrade pes position of claimed that Protochirotherium already appeared in the Late Lagerpeton chanarensis (according to Fechner, 2009). Scale = 1 cm, B without scale. Permian (Wuchiapingian). Until now Rotodactylus had not been
Fig.9. (A) Esqueleto pes de Lagerpeton chanarensis, (B) posição dos dígitos de pes do recorded from Biochron I. Klein and Lucas (2010) listed it as Lagerpeton chanarensis (de acordo com Fechner, 2009). Escala = 1 cm, B sem escala. first appearing in Biochron II, which is characterized by the presence of Chirotherium and corresponds to the late Olenekian Peabody (1948) reconstructed the producer of Rotodactylus to Anisian. The present study thus extends the known tracks as an agile reptile with dinosaur-like habitus. Haubold stratigraphic range of Rotodactylus. (1967) further elaborated on this idea. Haubold (1998) claimed that Inferences concerning the zoological affinities of the Rotodactylus is clearly a dinosauromorph ichnotaxon and thus producer of Rotodactylus are difficult due to the unusual position extends the known stratigraphic range of that group by several and orientation of digit V, the tip of which is always situated million years. Based on the same synapomorphies that they already posterior the impressions of the other four digits. Peabody (1948) used for interpreting Prorotodactylus Brusatte et al. (2010) also reconstructed a pes (Fig. 12 A) with a posteriorly rotated digit V considered Rotodactylus a dinosauromorph ichnogenus closely (see also Brusatte et al., 2010). Thulborn (1990) considered this related to Lagerpeton (see also Fechner, 2009 and Fig. 11). retroverted digit as a brace for the posterior part of the foot when However, the same concerns we registered about the interpretation it was lifted off the substrate. It is also possible that this digit was of Prorotodactylus also apply to that of Rotodactylus. spread to the side and backwards during digitigrade locomotion The Wolfhagen footprint assemblage only includes rare manus (Fig. 12 B), which would explain the slightly lateral position of "Dinosauromorph" tracks of Wolfhagen, Germany 87 the tip of the fifth digit in tracks from Wolfhagen. Ptaszyiński tracks represented an arboreal sidebranch of archosaurs, but there (2000) even suggested that the producer of the Rotodactylus exists no skeletal evidence for such a lineage.
Fig.11. Phylogenie of the Avemetatarsalia/Ornithodira according to Nesbitt (2011). The foot impressions of Prorotodactylus, Rotodactylus as well as the pedal skeleton of Lagerpeton are inserted in the scheme. Scale = 1 cm.
Fig.11. Filogenia dos Avemetatarsalia/Ornithodira de acordo com Nesbitt (2011). As impressões dos pés de Prorotodactylus, Rotodactylus, bem como dos esqueletos dos pés de Lagerpeton estão inseridas no esquema. Escala = 1 cm.
Fig.12. (A) digitigrade pes attitude and backward rotation of toe V of Rotodactylus ( reconstruction according to Peabody (1948). (B) stronger digitigrade pes attitude with an abducted toe V (not rotated backwards) of Rotodactylus. (C) digitigra de pes attitude with a more forward oriented toe V of Prorotodactylus. Scale = 1 cm.
Fig.12. (A) Atitude dos dígitos do pes e rotação traseira do dedo V de Rotodactylus (reconstrução de acordo com Peabody (1948). (B) Forte atitude dos dígitos do pes com o dedo V abductado (não rodado para trás) de Rotodactylus. (C) Atitude dos dígitos do pes com um dedo V mais orientado para a frente de Prorotodactylus. Escala = 1 cm.
6. Conclusions (Bad Hersfeld), Markus, Christian and Bastian Kunz (Wolfhagen) for the active help with the excavations in the quarry. We thank the From strata of the Detfurth sequence of the Middle Buntsandstein city of Wolfhagen as an owner of the quarry for the generous (Olenekian) exposed in a quarry near Wolfhagen (central kindness to let research proceed along the years. Germany) tracks of the "dinosauromorph" ichnogenera Prorotodactylus and Rotodactylus are described and References distinguished from the lepidosauromorph ichnotaxon Rhynchosauroides, which is very common in the same Avanzini, M., Bernardi, M., and Nicosia, U., 2011. The Permo-Triassic stratigraphic interval. Considerations concerning the pedal tetrapod faunal diversity in the Italian Southern Alps. In Ahmad Dar, I. morphology of "dinosauromorph" trackways lead to the and Ahmad Dar, M. (eds.), Earth and Environmental Sciences. 591- conclusion that the small dinosauromorph Lagerpeton is not 608. Brusatte, S.L., Niedzwiedzki, G., and Butler, R.J. 2010. Footprints pull related to the producer of such tracks contra Brusatte et al. origin and diversification of dinosaur stem lineage into Early Triassic. (2010). Thus the present study questions dinosauromorph Proceedings of the Royal Society, B 278, 1107-1113. affinities for the ichnogenus Prorotodactylus. The same also Demathieu, G., 1984. Une ichnofaune du Trias moyen du Bassin de appears to hold for Rotodactylus. Lodève (Herault, France). Annales de Paléontologie, 70, 47-273. The stratigraphically early occurrence of Rotodactylus as Demathieu, G., and Fichter, J., 1989. Die Karlshafener Fährten im well as the so far exclusive occurrence of Prorotodactylus in the Naturkundemuseum der Stadt Kassel – ihre Beschreibung und Germanic Basin suggest that the makers of these track types had Bedeutung. Phillipia, 6, 111-154. their origin and initial distribution in this region. Fechner, R., 2009. Morphofunctional Evolution of the Pelvic Girdle and Hindlimb of Dinosauromorpha on the Lineage to Sauropoda. Dissertation Universität München, 197. Acknowledgements Fichter, J., Heggemann, H., and Kunz, R., 1999. Neue bzw. bisher nicht veröffentlichte Tetrapodenfährten-Lokalitäten im Mittleren We are obliged to the following persons: Hans-Dieter Sues Buntsandstein Nordhessens und Südniedersachsens. Geologisches (Washington) for the translation into English, Klaus Hochberger Jahrbuch Hessen, 127, 33-55. 88 J. Fichter et al. / Comunicações Geológicas (2013) 100, 1, 81-88
Fichter, J., and Kunz, R., 2004. New genus and species of chirotherioid Klein, H., Voigt, S., Saber, H., Schneider, J.W., Hmninna, A., Fischer, J., tracks in The Detfurth Formation (Middle Bunter, Lower Triassic) of Lagnaoui, A., and Brosig, A., 2011. First occurrence of a Middle central Germany. Ichnos, 11, 183-193. Triassic tetrapod ichnofauna from the Argana Basin (Western High Fichter, J., and Kunz, R., 2006. Buntsandstein-Fossilien im Landkreis Atlas, Morocco). Palaeogeography, Palaeoclimatology, Kassel – Saurierspuren aus einer aufgelassenen Sandgrube im Palaeoecology, 307, 218-231. Stadtwald von Wolfhagen. Paläontologische Denkmäler in Hessen, 7, Kotanski, Z., Gierlinski, G., and Ptaszyiński, T., 2004. Reptile tracks 1-16. (Rotodactylus) from the Middle Triassic of the Djurdjura Mountains in Fichter, J., and Kunz, R., 2011. Neue Nachweise chirotherioider Fährten Algeria. Geological Quarterly (Warsaw), 48, 89-96. in der Detfurth-Formation (Mittlerer Buntsandstein, Untere Trias) bei Kunz, R., and Fichter, J., 2000. Die Wolfhager Saurierfährten. Wolfhagen. Geologisches Jahrbuch Hessen, 137, 5-18. Schriftenreihe des Vereins Regionalmuseum Wolfhagen, Reihe Fuglewicz, R., Ptaszyiński, T., and Rdzanek, K. 1990. Lower Triassic Forschungen, 9, 1-16. footprints from the Swietokrzyskie (Holy Cross) Mountains, Poland. Nesbitt, S.J., 2011. The early evolution of archosaurs: relationships and Acta Palaeontologica Polonica, 35, 109-164. the origin of major clades. Bulletin of the American Museum of Fortuny, J., Bolet, A., Sellés, A. G., Cartanyà, J., and Galobart, À., 2011. Natural History, 353, 1-292. New insights on the Permian and Triassic vertebrates from the Iberian Nopcsa, F., 1923. Die Familien der Reptilien. Fortschritte der Geologie Peninsula with emphasis on the Pyrenean and Catalonian basins. und Paläontologie, 2, 1-210. Journal of Iberian Geology, 37, 65-86 Peabody, F.E., 1948. Reptile and amphibian trackways from the Haubold, H., 1967. Eine Pseudosuchier-Fährtenfauna aus dem Moenkopi Formation of Arizona and Utah. University of California Buntsandstein Südthüringens. Hallesches Jahrbuch für Mitteldeutsche Publications, Bulletin of the Department of Geological Sciences, 27, Erdgeschichte, 8, 12-48. 295-468. Haubold, H., 1971a. Die Tetrapodenfährten des Buntsandsteins in der Ptaszyiński, T., 2000. Lower Triassic vertebrate footprints from Wióry, Deutschen Demokratischen Republik und in Westdeutschland und ihre Holy Cross Mountains, Poland. Acta Palaeontologica Polonica, 45, Äquivalente in der gesamten Trias. Paläontologische Abhandlungen, 151-194. A: Paläozoologie, 4, 395-548. Romer, A.S., 1971. The Chañares (Argentina) Triassic reptile fauna. X. Haubold, H., 1971b. Ichnia Amphibiorum et Reptiliorum fossilium. Two new but incompletely known long-limbed pseudosuchians. Handbuch der Paläoherpetologie, 18, 1-124. Breviora, 394, 1-7. Haubold, H., 1998. Tracks of the Dinosauromorpha from the Early Sereno, P.C., and Arcucci, A.B., 1994. Dinosaurian precursors from the Triassic. Hallesches Jahrbuch für Geowissenschaften, B, Beiheft, 5, Middle Triassic of Argentina: Lagerpeton chanarensis. Journal of 70-71. Vertebrate Paleontology, 13, 385-399. Haubold, H., 1999. Tracks of the Dinosauromorpha from the Lower Thulborn, R.A., 1990. Dinosaur Tracks. Chapman & Hall, London, 410. Triassic. Zentralblatt für Geologie und Paläontologie 1999, Teil I, Todesco, R., Wachtler, M., Kustatscher, E., and Avanzini, M., 2008. 783-795. Preliminary report on a new vertebrate track and flora site from Piz da Horn, M., 1976. Erläuterungen zur geologischen Karte von Hessen 1:25 Peres (Anisian-Illyrian): Olang Dolomites, Northern Italy. Geo.Alp, 5, 000, Blatt 4620 Arolsen. Geologisches Landesamt, Wiesbaden, 225. 121-137. Klein, H., and Haubold, H., 2007. Archosaur footprints – potential for Todesco, R., and Bernardi, M., 2011. Una nuova icnossociazione nel biochronology of Triassic continental sequences. In Lucas, S.G. and Triassico medio (Anisico) del Trentino meridionale (Val Gerlano, Spielmann, J.A. (eds.), The Global Triassic. New Mexico Museum of Vallarsa). Studi Trentini di Scienze Naturali, 88, 203-218. Natural History & Science Bulletin, 41, 120-130. Tourani, A., Benaouiss, N., Gand, G., Bourquinc, S., Jalil, N.-E., Broutin, Klein, H., and Lucas, S.G., 2010. Tetrapod footprints – their use in J., Battail, B., Germaine, D., Khaldoune, F., Sebban, S., Steyer, J.-S., biostratigraphy and biochronology of the Triassic. Geological Society and Vacante, R., 2010. Evidence of an Early Triassic age (Olenekian) of London Special Publikation, 334, 419-446 in Argana Basin (High Atlas, Morocco) based an new chirotherioid Klein, H., Voigt, S., Hminna, A., Saber, H., Schneider, J., and Hmich, D., traces. Comptes Rendus Palevol, 9, 201-208. 2010. Early Triassic archosaur-dominated footprint assemblage from the Argana Basin (Western High Atlas, Morocco). Ichnos, 17, 215- 227.