Amniota: Eureptilia) from the Upper Permian of Mallorca (Balearic Islands, Western Mediterranean)
Total Page:16
File Type:pdf, Size:1020Kb
A large multiple tooth-rowed captorhinid reptile (Amniota: Eureptilia) from the Upper Permian of Mallorca (Balearic Islands, Western Mediterranean) TORSTEN LIEBRECHT,1 JOSEP FORTUNY,2 ÀNGEL GALOBART,2 JOHANNES MÜLLER,1 and P. MARTIN SANDER,3 1 Museum für Naturkunde, Leibniz-Institut für Evolutions- und Biodiversitätsforschung, Invalidenstraße 43, 10115 Berlin, Germany, [email protected], [email protected]; 2 Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Edifici ICTA-ICP, Carrer de les Columnes s/n, Campus de la UAB, 08193 Cerdanyola del Vallès, Barcelona, Spain, [email protected], [email protected]; 3 Steinmann-Institut für Geologie, Mineralogie und Paläontologie, Rheinische Friedrich-Wilhelms-Universität Bonn, Nussallee 8, 53115 Bonn, Germany, [email protected] Supplementary data GEOLOGIC SETTING Today, the Balearic Islands (Spanish: Islas Baleares; Catalan: Illes Balears; Fig. S1) represent the geomorphologically highest, and thus emergent, parts of the north-eastern extension of the Betic Cordillera of southern Spain. This extension is also called the Balearic Promontory, which to the northwest is separated from the Iberian mainland by the Valencia Trough, an Oligocene to recent extensional structure that has a complex tectonic history closely connected to the Alpidic collisional movements that affected the western Mediterranean realm in the Late Mesozoic and Cenozoic (Roca, 1996). The northwestern part of the Island of Mallorca is occupied by the Serra de Tramuntana (Serra del Nord), a southwest-northeast trending horst-like structure that internally is built up of southeast-dipping Alpidic thrust sheets of unmetamorphosed, predominantly calcareous rocks of Jurassic age. Exposures of Permian to Middle Triassic terrestrial redbeds (so-called ‘Permo- Triassic’) are present only at the northwestern flank of the Serra de Tramuntana in the coastal area between the villages of Estellencs and Valldemosa. These deposits were referred to as ‘Buntsandstein facies’ (Rodríguez-Perea et al., 1987) and subdivided into three units which are named, from oldest to youngest, ‘Areniscas y Lutitas de Port des Canonge’, ‘Areniscas de Asá’, and ‘Lutitas y Areniscas de Son Serralta’ (Ramos, 1995). They represent the infill of post- variscan northwest-southeast trending extensional structures whose origin is closely related to that of the Permo-Triassic basins of the Iberian Chain (Gomez-Gras, 1993). The source area of the redbeds was the Hesperian Massif – the Iberian part of the Variscan mountains – to the northwest, and deposition took place under semi-arid conditions (Ramos, 1995; Linol et al., 2009). The relative age of the ‘Areniscas y Lutitas de Port des Canonge’ (for convenience hereafter referred to as Port des Canonge beds) is not well constrained and depends on whether its top is considered a major unconformity or not. Ramos and Doubinger (1989) identified a Late Permian (‘Thuringian’) palynoflora in the middle part of the overlying Asá beds and consequently also assigned the Port des Canonge beds to the Upper Permian. This was adopted 1 by most subsequent workers (Gomez-Gras, 1993; Ramos, 1995; Arche et al., 2002; Linol et al., 2009), except Bourquin et al. (2007) and Bourquin et al. (2011) who assume a major hiatus between the Port des Canonge and Asá beds and thus regard the former as Middle Permian. At least the Permian age now appears to be further corroborated by the occurrence of a captorhinid reptile. CIRCUMSTANCES OF DISCOVERY AND LOCAL GEOLOGY The maxilla was found incidentally by Mrs. Lieselotte Hannen, a hobby collector, on a beach of the Port des Canonge area (39° 41' 58" N, 2° 33' 04" E) in 2002. Port des Canonge is a small village in the municipality of Banyalbufar, about 15 km northwest of Palma de Mallorca. The beaches at Port des Canonge belong to the south-western part of the 80 km long north- western coast of Mallorca Island. The specimen was initially kept by the Hannen family and then, in 2007, was given to the Paleontological Department of the University of Bonn, Germany. From there it was transferred to the Museum für Naturkunde Berlin in 2008 for further study and preparation. The specimen is now accessioned at the Museu Balear de Ciències Naturals (MBCN) in Sóller, Island of Mallorca. The north-western coast of Mallorca is part of the north-western flank of the Serra de Tramuntana. It is a cliffed coast with many exposures of mainly Mesozoic rocks. The coastal cliffs at Port des Canonge (Fig. S1) are the type section of the deep red sandstones, siltstones, and claystones of the Port des Canonge beds, as discussed above. Because these siliciclastic redbeds much easier erode than the mainly calcareous rocks of the Serra de Tramuntana, the terrain in this area is generally lower and the slopes are less steep than in the adjacent areas. Small ephemeral streams (locally referred to as ‘torrentes’) have cut several small valleys into the ground that open into the beach at their lower end. According to Mrs. Hannen the maxilla was found among the pebbles of the beach gravel at Cala Gata, a few hundred meters W of Port des Canonge (Figs. 2S, 3S). The gravel forms a thin veneer on Permian mudstones, which also are exposed in the coastal cliff, and finer grained Quaternary deposits of reworked Permian redbeds. Closer examinations of these underlying deposits did not lead to discoveries of further bone material. The discovery in the gravel indicates that the specimen must have been transported, but only a very short distance as shown by the lack of abrasion. The sediment adhering to the specimen allows an assignment to the siltstones and mudstones exposed in the coastal section of the Port des Canonge area. Hence, the maxilla must either have immediately been weathered out of the coastal cliff or transported via an ephemeral stream over a very short distance from an inland exposure of the same lithologic unit (i.e. the Port des Canonge beds). The latter option is supported by the fact that the exact site of discovery is situated immediately below the exit of the valley of the Torrent son Bunyola. Our field work confirmed that the siliciclastic redbeds of the Port des Canonge area are clearly terrestrial. They show features of fluvial deposition (channel and overbank flood deposits) as well as of soil formation such as root traces. 2 FIGURE S1. Map showing the geographic position and geological setting of Cala Gata west-northwest of the village of Port des Canonge, the site where MBCN 15730 has been found. FIGURE S2. Coastal exposures and beach of Cala Gata west-northwest of Port des Canonge, seen from NE. Permian redbeds form the gently rolling reddish hills down at the coast line. The mountain tops and the prominent headland further in the back (Punta de s’Àguila) are made up of Mesozoic and Cenozoic carbonate rocks. MBCN 15730 was found on the gravel beach in the foreground. 3 FIGURE S3. Satellite image of the coastal area immediately west of Port des Canonge, Illes Balears. The place where the specimen was encountered in 2002 is marked by the circle. PRESERVATION MBCN 15730 was preserved in a matrix of red siltstone. The original bone tissue is reddish-brown due to impregnation with or replacement by iron minerals. After preparation most of the original bone surface is easily identifiable by a cover of a light greyish-blue secondary mineral, probably the phosphate mineral vivianite. CLADISTIC ANALYSIS We used a modified dataset of Reisz et al. (2015), which is the most recent version of a dataset used and successively expanded in earlier works (Dodick and Modesto, 1995; Müller and Reisz, 2005, 2006; Modesto et al., 2007; Sumida et al., 2010; Reisz et al., 2011; Modesto et al., 2014). Only seven of 75 characters could be coded (chars. no. 6, 7, 9, 10, 12, 20, 39; see supplementary material) for MBCN 15730. Almost all are dental characters. The analysis was conducted using the ‘implicit enumeration’ algorithm of TNT (Goloboff et al., 2008), which is essentially the same as the ‘branch-and-bound’ algorithm in PAUP. All characters were left unordered and unweighted. The ‘protorothyridid’ eureptile Protorothyris was selected as outgroup. Tree statistics and support values were obtained with the script statss.run and implemented features in TNT. Coding of characters for the Mallorca specimen (MBCN 15730) as added to the data matrix of Reisz et al. (2015): Mallorca specimen ?????31?13 ?2???????0 ?????????? ????????1? ?????????? ?????????? ?????????? ????? 4 FIGURE S4. Strict consensus of the two trees obtained from the TNT analysis of the dataset of Reisz et al. (2015). Numbers on nodes refer to bootstrap values (above) and bremer (decay) indices (below). LITERATURE CITED Arche, A., J. López-Gómez, and H. Vargas. 2002. Propuesta de correlación entre los sedimentos Pérmicos y Triásicos de la Cordillera Ibérica Este y de las Islas Baleares. Geogaceta 32:275-278 Bourquin, S., A. Bercovici, J. López-Gómez, J. B. Diez, J. Broutin, A. Ronchi, M. Durand, A. Arché, B. Linol, and F. Amour. 2011. The Permian–Triassic transition and the onset of Mesozoic sedimentation at the northwestern peri-Tethyan domain scale: Palaeogeographic maps and geodynamic implications. Palaeogeography, Palaeoclimatology, Palaeoecology 299:265-280 Bourquin, S., M. Durand, J. B. Diez, J. Broutin, and F. Fluteau. 2007. The Permian-Triassic boundary and Lower Triassic sedimentation in western European basins: an overview. Journal of Iberian Geology 33:221-236 Dodick, J. T., and S. P. Modesto. 1995. The cranial anatomy of the captorhinid reptile Labidosaurikos meachami from the Lower Permian of Oklahoma. Palaeontology 38:687-711 Goloboff, P. A., J. S. Farris, and K. C. Nixon. 2008. TNT, a free program for phylogenetic analysis. Cladistics 24:774-786 5 Gomez-Gras, D. 1993. El Permotrías de las Baleares y de la vertiente mediterránea de la Cordillera Ibérica y del Maestrat: Facies y Petrología Sedimentaria (Parte II).