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Journal of Vertebrate Paleontology a New Species of Metoposaurus From This article was downloaded by: [Faculdade de Ciencias Sociais e Humanas] On: 24 March 2015, At: 02:17 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK Journal of Vertebrate Paleontology Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/ujvp20 A new species of Metoposaurus from the Late Triassic of Portugal and comments on the systematics and biogeography of metoposaurid temnospondyls Stephen L. Brusattea, Richard J. Butlerbc, Octávio Mateusde & J. Sébastien Steyerf a School of GeoSciences, University of Edinburgh, Grant Institute, The King's Buildings, James Hutton Road, Edinburgh, EH9 3FE, U.K. b School of Geography, Earth and Environmental Sciences, University of Birmingham, Edgbaston, Birmingham B15 2TT, U.K. c GeoBio-Center, Ludwig-Maximilians-Universität München, Richard-Wagner-Str. 10, 80333 Munich, Germany; Click for updates d GeoBioTec-Faculdade de Ciências e Tecnologia, Universidade Nova de Lisboa, 2829-516 Caparica, Portugal e Museu da Lourinhã, 2530-157, Lourinha, Portugal f CNRS UMR 7207, Centre de Recherches en Paléobiodiversité et Paléoenvironnements, CNRS- MNHN-UPMC, 8 rue Buffon, CP38, 75005 Paris, France Published online: 23 Mar 2015. To cite this article: Stephen L. Brusatte, Richard J. Butler, Octávio Mateus & J. Sébastien Steyer (2015): A new species of Metoposaurus from the Late Triassic of Portugal and comments on the systematics and biogeography of metoposaurid temnospondyls, Journal of Vertebrate Paleontology, DOI: 10.1080/02724634.2014.912988 To link to this article: http://dx.doi.org/10.1080/02724634.2014.912988 PLEASE SCROLL DOWN FOR ARTICLE Taylor & Francis makes every effort to ensure the accuracy of all the information (the “Content”) contained in the publications on our platform. However, Taylor & Francis, our agents, and our licensors make no representations or warranties whatsoever as to the accuracy, completeness, or suitability for any purpose of the Content. Any opinions and views expressed in this publication are the opinions and views of the authors, and are not the views of or endorsed by Taylor & Francis. The accuracy of the Content should not be relied upon and should be independently verified with primary sources of information. Taylor and Francis shall not be liable for any losses, actions, claims, proceedings, demands, costs, expenses, damages, and other liabilities whatsoever or howsoever caused arising directly or indirectly in connection with, in relation to or arising out of the use of the Content. This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. Terms & Conditions of access and use can be found at http:// www.tandfonline.com/page/terms-and-conditions Journal of Vertebrate Paleontology e912988 (23 pages) Ó by the Society of Vertebrate Paleontology DOI: 10.1080/02724634.2014.912988 ARTICLE A NEW SPECIES OF METOPOSAURUS FROM THE LATE TRIASSIC OF PORTUGAL AND COMMENTS ON THE SYSTEMATICS AND BIOGEOGRAPHY OF METOPOSAURID TEMNOSPONDYLS STEPHEN L. BRUSATTE,*,1 RICHARD J. BUTLER,2,3 OCTAVIO MATEUS,4,5 and J. SEBASTIEN STEYER6 1School of GeoSciences, University of Edinburgh, Grant Institute, The King’s Buildings, James Hutton Road, Edinburgh, EH9 3FE, U.K., [email protected]; 2School of Geography, Earth and Environmental Sciences, University of Birmingham, Edgbaston, Birmingham B15 2TT, U.K., [email protected]; 3GeoBio-Center, Ludwig-Maximilians-Universitat€ Munchen,€ Richard-Wagner-Str. 10, 80333 Munich, Germany; 4GeoBioTec-Faculdade de Ciencias^ e Tecnologia, Universidade Nova de Lisboa, 2829-516 Caparica, Portugal, [email protected]; 5Museu da Lourinha,~ 2530-157, Lourinha, Portugal 6CNRS UMR 7207, Centre de Recherches en Paleobiodiversit e et Paleoenvironnements, CNRS-MNHN-UPMC, 8 rue Buffon, CP38, 75005 Paris, France, [email protected] ABSTRACT—Metoposaurids are a group of temnospondyl amphibians that filled crocodile-like predatory niches in fluvial and lacustrine environments during the Late Triassic. Metoposaurids are common in the Upper Triassic sediments of North Africa, Europe, India, and North America, but many questions about their systematics and phylogeny remain unresolved. We here erect Metoposaurus algarvensis, sp. nov., the first Metoposaurus species from the Iberian Peninsula, based on several new specimens from a Late Triassic bonebed in Algarve, southern Portugal. We describe the cranial and pectoral anatomy of M. algarvensis and compare it with other metoposaurids (particularly other specimens of Metoposaurus from Germany and Poland). We provide a revised diagnosis and species-level taxonomy for the genus Metoposaurus, which is currently represented with certainty by three European species (M. diagnosticus, M. krasiejowensis, M. algarvensis). We also identify cranial characters that differentiate these three species, and may have phylogenetic significance. These include features of the braincase and mandible, which indicate that metoposaurid skulls are more variable than previously thought. The new Portuguese bonebed provides further evidence that metoposaurids congregated in fluvial and lacustrine settings across their geographic range and often succumbed to mass death events. We provide an updated paleogeographic map depicting all known metoposaurid occurrences, which shows that these temnospondyls were globally distributed in low latitudes during the Late Triassic and had a similar, but not identical, paleogeographic range as phytosaurs. http://zoobank.org/urn:lsid:zoobank.org:pub:083C80C6-0AB6-49E1-A636-6A8BDBC06A47 INTRODUCTION (1.25–3 m body length), and filled crocodile-like predatory niches in lacustrine and fluvial environments (e.g., Hunt, 1993; The Late Triassic (Carnian–Rhaetian, ca. 237–201 million Schoch and Milner, 2000). years ago) was a formative phase in the evolution of ‘modern’ Metoposaurids are some of the most common and character- terrestrial ecosystems dominated by familiar clades such as mam- istic non-marine vertebrates from the Late Triassic, and their mals, lizards, turtles, and archosaurs (e.g., Sues and Fraser, distinctly ornamented skulls, spool-shaped vertebrae, and atro- 2010). By this time, ecosystems had largely stabilized after the phied limb elements are found at numerous sites in Africa (e.g., unprecedented devastation of the end-Permian extinction (e.g., Dutuit, 1976), Europe (e.g., Meyer, 1842; Fraas, 1913; Kuhn, Chen and Benton, 2012) and soon-to-be dominant groups such Downloaded by [Faculdade de Ciencias Sociais e Humanas] at 02:17 24 March 2015 1933; Sulej, 2002, 2007; Milner and Schoch, 2004), India (e.g., as dinosaurs were experiencing their initial radiations (e.g., Bru- Chowdhury, 1965; Sengupta, 1992), and North America (e.g., satte et al., 2010; Langer et al., 2010; Sidor et al., 2013). Case, 1931; Sawin, 1945; Romer, 1947; Colbert and Imbrie, Although faunas were clearly ‘modernizing’ during the Late Tri- 1956; Hunt, 1993; Long and Murry, 1995). Metoposaurid speci- assic, some ‘archaic’ groups of amphibians, reptiles, and synap- mens from the Germanic Basin of Europe were some of the sids remained diverse and abundant, with some undergoing their first temnospondyls to be described, and in the 150+ years since final bursts of evolutionary radiation before being ravaged by these original materials were studied hundreds of new speci- another mass extinction at the end of the Triassic. Among the mens from Germany and Poland have helped shed light on the most prominent of these groups is the Metoposauridae, a clade paleoecology (Bodzioch and Kowal-Linka, 2012), intraspecific of temnospondyl amphibians that were globally distributed in and ontogenetic variation (Sulej, 2007), bone histology low paleolatitudes during the Late Triassic (particularly during (Konietzko-Meier and Klein, 2013), and growth (Sulej, 2007; the early Carnian–middle Norian), achieved mid- to large size Konietzko-Meier and Klein, 2013) of extinct amphibians. Despite these advances in understanding metoposaurid biology, however, many fundamental questions about metoposaurid sys- *Corresponding author. tematics and phylogeny remain unresolved (e.g., Hunt, 1993; Color versions of one or more of the figures in this article can be found Schoch and Milner, 2000; Sulej, 2002, 2007). Perhaps most online at www.tandfonline.com/ujvp. e912988-2 JOURNAL OF VERTEBRATE PALEONTOLOGY FIGURE 1. A, regional stratigraphic section of the Triassic sedimentary rocks of southern Portugal; B, location map of the Algarve Basin of southern Portugal, with the Triassic exposures shaded gray. problematic, there is still no consensus on the alpha-level taxon- cladistic analyses. We also comment on the biogeographic distri- omy of the group, largely because numerous species have been bution of metoposaurids, provide an updated paleogeographic named over the past century and a half, often based on frag- map denoting all known metoposaurid occurrences, and discuss mentary material and without a detailed understanding of intra- how the Portuguese specimens lend further credence to the specific variation. widespread distribution of these amphibians in fluvial and lacus- We here describe new well-preserved specimens of the meto- trine environments in low paleolatitudes during the Late posaurid
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