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Journal of Vertebrate Paleontology Cranial Anatomy, Ontogeny, And This article was downloaded by: [Universite Laval] On: 09 July 2014, At: 04:25 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 Cranial anatomy, ontogeny, and relationships of the Late Carboniferous tetrapod Gephyrostegus bohemicus Jaekel, 1902 Jozef Klembaraa, Jennifer A. Clackb, Andrew R. Milnerc & Marcello Rutad a Department of Ecology, Faculty of Natural Sciences, Comenius University in Bratislava, Mlynská dolina, 84215 Bratislava, Slovakia b University Museum of Zoology, Downing Street, Cambridge, CB2 3EJ, U.K. c Department of Earth Sciences, The Natural History Museum, Cromwell Road, London SW7 5BD, U.K. d School of Life Sciences, University of Lincoln, Riseholme Park, Lincoln LN2 2LG, U.K. Published online: 08 Jul 2014. To cite this article: Jozef Klembara, Jennifer A. Clack, Andrew R. Milner & Marcello Ruta (2014) Cranial anatomy, ontogeny, and relationships of the Late Carboniferous tetrapod Gephyrostegus bohemicus Jaekel, 1902, Journal of Vertebrate Paleontology, 34:4, 774-792 To link to this article: http://dx.doi.org/10.1080/02724634.2014.837055 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. 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Terms & Conditions of access and use can be found at http:// www.tandfonline.com/page/terms-and-conditions Journal of Vertebrate Paleontology 34(4):774–792, July 2014 © 2014 by the Society of Vertebrate Paleontology ARTICLE CRANIAL ANATOMY, ONTOGENY, AND RELATIONSHIPS OF THE LATE CARBONIFEROUS TETRAPOD GEPHYROSTEGUS BOHEMICUS JAEKEL, 1902 JOZEF KLEMBARA,*,1 JENNIFERA.CLACK,2 ANDREW R. MILNER,3 and MARCELLO RUTA4 1Department of Ecology, Faculty of Natural Sciences, Comenius University in Bratislava, Mlynska´ dolina, 84215 Bratislava, Slovakia, [email protected]; 2University Museum of Zoology, Downing Street, Cambridge CB2 3EJ, U.K., [email protected]; 3Department of Earth Sciences, The Natural History Museum, Cromwell Road, London SW7 5BD, U.K., [email protected]; 4School of Life Sciences, University of Lincoln, Riseholme Park, Lincoln LN2 2LG, U.K., [email protected] ABSTRACT—We review the cranial morphology of the Late Carboniferous terrestrial tetrapod Gephyrostegus bohemicus from the coal deposits of the N´yranyˇ Basin in the Czech Republic. Gephyrostegus is known from several skulls ranging in length from about 25 mm to about 58 mm (holotype). The narrow skull is about twice as long as wide and shows a well-ossified quadrate and articular, but no evidence of braincase ossification. Autapomorphic features include a pustular ornamentation on some skull table bones, and a plate-like tabular process exhibiting a fine dorsal pitting. Gephyrostegus shares with Brukter- erpeton fiebigi (Late Carboniferous, Germany) the presence of low, anteromedially to posterolaterally orientated sharp ridges on the posteroventral surface of the vomer. It shares with seymouriamorphs a rectangular, transverse pterygoid process and closely packed, radially arranged rows of small denticles on the palate. A phylogenetic analysis retrieves Gephyrostegidae (Gephyrostegus, Bruktererpeton) as sister group to Seymouriamorpha, although this wider clade receives low bootstrap sup- port. SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP INTRODUCTION each other. We have been able to study additional specimens unavailable to Carroll, and unrecognized until recently. Our re- A wealth of tetrapod taxa, most with body lengths of around description aims to reconcile the conflicting skull reconstructions 140 mm, have been described from N´yranyˇ and Tremoˇ snˇ a(Plze´ n-ˇ in Carroll and to provide a new cladistic analysis incorporating Manetˇ ın´ Basin, Czech Republic). Among them, the genus Gephy- our new observations on the cranial morphology of Gephyroste- rostegus was last described in detail by Carroll (1970), with gus and of a range of other taxa not available to Carroll. Our some amendments by Panchen (1970). According to Carroll cladistic analysis also includes characters from the postcranium. (1970:305), Gephyrostegus “represents the morphological pattern However, these will be detailed in a separate work on the expected in the group of anthracosaurs which gave rise to rep- postcranial osteology of the Carboniferous Czech anthracosaurs. tiles,” and forms a branch of the anthracosaurs separate from em- Below, we provide a detailed historical background to research Downloaded by [Universite Laval] at 04:25 09 July 2014 bolomeres (here, ‘anthracosaur’ refers to the group that includes on this faunistic assemblage. eoherpetontids and embolomeres; Holmes, 1984, 1989; Smithson, 1985; Panchen and Smithson, 1988; Ruta and Clack, 2006; Ruta and Coates, 2007). Its robust postcranium and gracile limbs sug- HISTORICAL BACKGROUND gest a terrestrial animal, in contrast to the largely aquatic em- bolomeres. Although Carroll’s conclusions on the affinities of this Since their first description (Fritsch, 1885a, 1885b), the small taxon have been superseded by cladistic studies, in many of these anthracosaurs from N´yranyˇ and Tremoˇ snˇ a´ have often been in- Gephyrostegus has appeared as a stem amniote, often crownward voked in relation to the origin of amniotes and other amniote- of embolomeres (e.g., Clack and Finney, 2005; Ruta and Clack, like groups, such as seymouriamorphs (e.g., Panchen and Smith- 2006; Ruta and Coates, 2007). son, 1988; Lee and Spencer, 1997). The specimens are dispersed Given the significance ascribed to this genus, it is unfortunate across several museums, and the political history of the 20th cen- that Carroll was unable to complete his studies (see below) and tury has meant that they were not all readily accessible to most had to rely on only two specimens rendered as peels. He also potential researchers. Most previous studies have been based on included descriptions and figures of some of the other N´yranyˇ one to four specimens and no author has attempted to incorpo- tetrapods: Diplovertebron punctatum, Eusauropleura digitata, rate all of the determinate material in a single description. The and Solenodonsaurus janenschi. These are dealt with briefly be- material discussed here has a complicated history, having been low, but are not otherwise described. Carroll provided detailed described as five different species in four different genera. One drawings of Gephyrostegus, a full skeletal reconstruction, and publication (Brough and Brough, 1967) added some basal am- two alternative skull reconstructions based on the two speci- niotes to the hypodigm, but these were later removed (Carroll, mens studied. As a consequence, they showed differences from 1970). Because the full taxonomic and descriptive history of this material has never been reviewed, we take the opportunity to outline it here. The material originally described by Antonın´ Fricˇ *Corresponding author. (or Anton Fritsch), referred to by his Fritsch Original (Fr. Orig.) 774 KLEMBARA ET AL.—CRANIAL ANATOMY OF GEPHYROSTEGUS BOHEMICUS 775 numbers, is held in the National Museum, Prague, and now has specimen from N´yranyˇ (DMSW B.65) for his own collection. NMP catalog numbers. He concluded that not only was it a small example of Jaekel’s Fritsch had collected some of this material from the mine at Gephyrostegus, but also that both were the same as Fritsch’s first- N´yranyˇ in the 1870s, but did not mention it in his first report on described anthracosaur, Diplovertebron punctatum.Datafrom the fauna (Fric,ˇ 1876). By the time he had completed the first these specimens allowed Watson (1926:figs. 29–31) to provide a part of his monumental ‘Fauna der Gaskohle und der Kalksteine fairly complete reconstruction of Diplovertebron punctatum (= der Permformation Bohmens’¨ (Fritsch, 1879), he had recognized Gephyrostegus bohemicus)asasmallN´yranyˇ anthracosaur. some of the vertebrate material as distinctive and gave advanced In the 1930s, Watson’s research assistant, Margaret Steen, notice of the taxon Diplovertebron punctatum diagnosed by its published several papers dealing with Permo-Carboniferous am- double centra. This and other taxa listed on pages 26–32 of the phibians. In one of these, Steen (1938) reviewed the N´yranyˇ first part of ‘Fauna
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