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( Metoposaurus) Azerouali (Dutuit) Comb Redescription of Arganasaurus ( Metoposaurus) azerouali (Dutuit) comb. nov. from the Upper Triassic of the Argana Basin (Morocco), and the first phylogenetic analysis of the Metoposauridae (Amphibia, Temnospondyli) Valentin Buffa, Nour-Eddine Jalil, J.-Sébastien Steyer To cite this version: Valentin Buffa, Nour-Eddine Jalil, J.-Sébastien Steyer. Redescription of Arganasaurus (Meto- posaurus) azerouali (Dutuit) comb. nov. from the Upper Triassic of the Argana Basin (Morocco), and the first phylogenetic analysis of the Metoposauridae (Amphibia, Temnospondyli). Special pa- pers in palaeontology, Wiley Blackwell Publishing, 2019, 5 (4), pp.699-717. 10.1002/spp2.1259. hal-02967815 HAL Id: hal-02967815 https://hal.sorbonne-universite.fr/hal-02967815 Submitted on 15 Oct 2020 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Page 1 of 60 Palaeontology 1 2 3 REDESCRIPTION OF ARGANASAURUS (METOPOSAURUS) AZEROUALI (DUTUIT) 4 5 6 COMB. NOV. FROM THE LATE TRIASSIC OF THE ARGANA BASIN (MOROCCO), 7 8 AND THE FIRST PHYLOGENETIC ANALYSIS OF THE METOPOSAURIDAE 9 10 (AMPHIBIA, TEMNOSPONDYLI) 11 12 13 14 15 16 by VALENTIN BUFFA*1, NOUR-EDDINE JALIL1,2 and J.-SEBASTIEN STEYER1 17 18 19 1Centre de Recherche sur la paléobiodiversité et les paléoenvironnements, UMR 7207 CNRS- 20 21 MNHN-UPMC, Muséum national d’Histoire naturelle, CP38, 8 rue Buffon, 75005 Paris ; e- 22 23 24 mails: [email protected], [email protected], jean- 25 26 [email protected] 27 28 29 2BioDEcos Département de géologie, Faculté des Sciences Semlalia, Université Cadi Ayyad, 30 31 Marrakech, Maroc 32 33 34 *Corresponding author 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 1 Palaeontology Palaeontology Page 2 of 60 1 2 3 Abstract: A systematic revision of the temnospondyl “Metoposaurus” azerouali Dutuit from 4 5 6 the Late Triassic of the Argana Basin (Western High Atlas, Morocco) is presented. The type 7 8 material is redescribed in detail, and a preliminary phylogenetic analysis - the first one dealing 9 10 with all metoposaurid species - is also conducted in order to test its position within the 11 12 Metoposauridae. Our analysis places “Metoposaurus” azerouali as sister-taxon to 13 14 15 Arganasaurus lyazidi in a robust clade supported by two unambiguous synapomorphies 16 17 (buldge-like tabular horn and exoccipital process visible in dorsal view) and two ambiguous 18 19 synapomorphies (cultriform process of uniform width and subtriangular posterior Meckelian 20 21 22 fenestra). We therefore propose the new combination Arganasaurus azerouali comb. nov. for 23 24 the species “M.” azerouali. Our analysis also confirms that the central Laurasian genus 25 26 Metoposaurus is monophyletic but not the genus Koskinonodon which deserves its own 27 28 systematic revision. By consequent, the Late Triassic rich vertebrate fauna from the Argana 29 30 31 Basin comprises the metoposauroid Almasaurus habazzi, the basal metoposaurid Dutuitosaurus 32 33 ouazzoui, and the genus Arganasaurus which is represented by the type species A. lyazidi and 34 35 A. azerouali comb. nov. Combined with the stratigraphic and geographic occurrences of the 36 37 38 taxa, our phylogenetic analysis suggests that metoposaurids may have appeared during the 39 40 Longobardian (late Ladinian) in central Pangea. Their diversification may be linked to the 41 42 Carnian Pluvial Episode. At least, their extinction may have occurred during the Rhaetian 43 44 45 because of the aridification of the climate and/or competition with amniotes. 46 47 48 Keywords: phylogenetic analysis, Metoposauridae, diversification, Carnian Pluvial Episode, 49 50 temnospondyl, Morocco. 51 52 53 54 55 56 The Argana Basin, also called the Argana Corridor (Western High Atlas, Morocco), provides a 57 58 series of continental deposits spanning from the Middle-Late Permian to the early Jurassic. 59 60 Since the 1960s, this basin has yielded numerous fossils that document a key period in the 2 Palaeontology Page 3 of 60 Palaeontology 1 2 3 evolutionary history of terrestrial vertebrates, including the Permo-Triassic mass extinction 4 5 6 (Khaldoune et al. 2017). 7 8 Among the Triassic amphibians recovered from the Argana Basin, three species of 9 10 11 metoposaurids are known: Dutuitosaurus ouazzoui (Dutuit, 1976), Arganasaurus lyazidi 12 13 (Dutuit, 1976) and “Metoposaurus” azerouali Dutuit, 1976. The latter has been described 14 15 briefly in French by Dutuit (1976, p. 165-176) and its systematic status is controversial as it has 16 17 18 been considered as a nomen dubium by Hunt (1993) but a valid taxon by Schoch and Milner 19 20 (2000). 21 22 23 The family Metoposauridae Watson, 1919 includes temnospondyl amphibians which 24 25 occupied freshwater predatory niches during the Late Triassic, especially between the middle 26 27 28 Carnian and the Norian (Hunt 1993; Schoch and Milner 2000; Sulej 2002; Brusatte et al. 2015). 29 30 Metoposaurids are found in low palaeolatitude deposits in Europe, India, Morocco and North 31 32 America (Case 1922, 1931; Dutuit 1976; Hunt 1993; Sengupta 2002; Sulej 2002; Brusatte et 33 34 al. 2015) and presumably in Algeria, France and Madagascar (Schoch and Milner 2000). The 35 36 37 position of the Metoposauridae within temnospondyls has been an ongoing debate for decades, 38 39 but the current consensus considers them as trematosaurian stereospondyls (Schoch and Milner 40 41 2000; Yates and Warren 2000; Damiani and Yates 2003; Schoch 2008, 2011, 2013). 42 43 44 Several phylogenetic studies involving metoposaurids have been done (Hunt 1993; 45 46 47 Steyer 2002; Ruta et al. 2007; Sulej 2007; Schoch 2008, 2011, 2013; McHugh 2012) but the 48 49 only computer-based phylogenetical analyses to date were conducted on temnospondyls or 50 51 stereospondyls as a whole and did not take into account all metoposaurid species. This study 52 53 provides a systematic revision of the species “Metoposaurus” azerouali Dutuit, 1976 and the 54 55 56 first preliminary phylogenetic analysis of the family Metoposauridae. 57 58 59 60 3 Palaeontology Palaeontology Page 4 of 60 1 2 3 Institutional abbreviation. BMNH, British Museum of Natural History, London; MNHN, 4 5 6 Muséum national d’Histoire naturelle, Paris. 7 8 9 10 11 SYSTEMATIC PALAEONTOLOGY 12 13 14 Order TEMNOSPONDYLI Zittel, 1888 15 16 17 Suborder STEREOSPONDYLI Zittel, 1888 18 19 20 Family METOPOSAURIDAE Watson, 1919 21 22 23 Phylogenetic definition. Node-based clade including the last common ancestor of 24 25 26 Dutuitosaurus ouazzoui and Metoposaurus diagnosticus (Meyer, 1842) and all of its 27 28 descendants (see analysis below). 29 30 31 Diagnosis (modified from Sulej 2007). Large oval and widely separated orbits in the anterior 32 33 half of the skull; parasphenoid central depression; broad cultriform process; absence of 34 35 cultriform process keel; quadrate excluded from squamosal by quadratojugal; quadrate condyle 36 37 38 subtriangular in ventral view; paraquadrate foramen present; deep and rounded sinus 39 40 pterygoidei below the oblique crest on the posterior side of the ascending ramus of the 41 42 pterygoid; mandibular dentition strictly on the dentary; prearticular not anteriorly positioned to 43 44 45 the posterior coronoid; articular extending posteriorly between the surangular and the 46 47 prearticular along the dorsal surface of the retroarticular process; parapophysis on two 48 49 neighboring intercentra of presacral and sacral vertebra; neural spine dorsally extended in 50 51 52 posterior trunk or tail; clavicular blade indentation; clavicular contact above the interclavicle; 53 54 unusually large manus and pes; concave anterior margin of the ilium shaft; main axes of the 55 56 proximal and distal heads of the femur set to each other at 90°. 57 58 59 60 4 Palaeontology Page 5 of 60 Palaeontology 1 2 3 Content. The type species Metoposaurus diagnosticus from Germany; Metoposaurus 4 5 6 krasiejowensis (Sulej, 2002) from Poland (we followed Brusatte et al. 2015 in considering the 7 8 subspecies “M. diagnosticus krasiejowensis” Sulej, 2002 as the species M. krasiejowensis); 9 10 Metoposaurus algarvensis Brusatte, Butler, Mateus and Steyer, 2015 from Portugal; 11 12 Koskinonodon perfectus (Case, 1922) and Koskinonodon bakeri (Case, 1931) (following the 13 14 15 generic attribution of Sulej 2007 and Brusatte et al. 2015) from North America; Koskinonodon 16 17 maleriensis (Chowdhury, 1965) from India (following the generic attribution of Sengupta 2002, 18 19 2003 and the remarks of Brusatte et al. 2015); Dutuitosaurus ouazzoui, Arganasaurus lyazidi 20 21 22 and “Metoposaurus” azerouali from Morocco. 23 24 25 Remarks. In his diagnosis of the Metoposauridae, Sulej (2007) also added large external nostrils 26 27 and an exoccipital-pterygoid suture perpendicular to the skull axis in ventral view. However, 28 29 as these characters are also shared with Callistomordax kugleri Schoch, 2008, sister-taxon to 30 31 the Metoposauridae in our analysis (see below), we consider them as no longer valid for this 32 33 34 diagnosis. 35 36 37 In his revision, Hunt (1993) discriminated the genera Koskinonodon (Case, 1922) and 38 39 Metoposaurus Lydekker, 1890 on the basis of the lacrimal entering or not the orbit 40 41 (respectively). However, as recent works showed that the lacrimal does enter the orbit in the 42 43 44 genus Metoposaurus (Sulej 2002, 2007; Brusatte et al. 2015), this discrimination is no longer 45 46 valid. While the diagnosis of the genus Metoposaurus has been amended (Sulej 2002, 2007; 47 48 Brusatte et al.
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