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Pleurodira) Identified in the British Record: Palemys Bowerbankii New Combination

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Pleurodira) Identified in the British Record: Palemys Bowerbankii New Combination Palaeontologia Electronica palaeo-electronica.org New insights on the only bothremydid turtle (Pleurodira) identified in the British record: Palemys bowerbankii new combination Adán Pérez-García ABSTRACT The British record of Pleurodira is very limited compared to that of Continental Europe. It is restricted to two lower Eocene coastal forms, corresponding to two lin- eages of Pelomedusoides: Podocnemididae and Bothremydidae. One of them is a recently recognized indeterminate species of the podocnemidid Eocenochelus, repre- sented by a single individual from Bracklesham Bay (Sussex). Eocenochelus is a lin- eage of African origin that reached Europe during the Paleogene and is recognized in several European countries. The other British form is represented by several speci- mens from the London Clay of the Isle of Sheppey (Kent). Several shells were identi- fied there, since the 19th century, allowing the description of four putative new species. The detailed study of the classic and subsequently found specimens, including unpub- lished individuals, confirms their attribution to a single species, for which the new com- bination Palemys bowerbankii is proposed. It is the only pleurodiran turtle recognized in the British record at a specific level. A neotype is designated, and a new diagnosis is provided. Photographs of most of the classic specimens of Palemys bowerbankii are included here for the first time. The detailed study of all these specimens improves the knowledge of this species. With this information, this exclusively British form is included for the first time in a cladistic analysis. It is not interpreted as the result of a Cenozoic dispersal event from Africa, as was the case with Eocenochelus, but as a member of a well-established lineage present in Europe during the Late Cretaceous, which lived in this continent at least until the early Eocene. Adán Pérez-García. Grupo de Biología Evolutiva, Facultad de Ciencias, UNED, Paseo Senda del Rey, 9, 28040 Madrid, Spain. [email protected] Keywords: Pelomedusoides; Bothremydidae; early Eocene; London Clay Submission: 11 January 2018 Acceptance: 2 July 2018 Pérez-García, Adán. 2018. New insights on the only bothremydid turtle (Pleurodira) identified in the British record: Palemys bowerbankii new combination. Palaeontologia Electronica 21.2.28A 1-12. https://doi.org/10.26879/849 palaeo-electronica.org/content/2018/2268-british-bothremydid-turtle Copyright: August 2018 Paleontology Society This is an open access article distributed under the terms of Attribution-NonCommercial-ShareAlike 4.0 International (CC BY-NC-SA 4.0), which permits users to copy and redistribute the material in any medium or format, provided it is not used for commercial purposes and the original author and source are credited, with indications if any changes are made. creativecommons.org/licenses/by-nc-sa/4.0/ PÉREZ-GARCÍA: BRITISH BOTHREMYDID TURTLE INTRODUCTION these classic specimens had been previously pub- lished. With the exception of its holotype, which is The British turtle record includes several identified as lost since the 19th century (Lydekker Mesozoic members of the stem group of Testu- and Boulenger, 1887), photographs of all the clas- dines, as well as a great diversity of representa- sic specimens are included here for the first time. tives of Cryptodira, both from Mesozoic and In addition, shells subsequently found, including Cenozoic horizons (Lapparent de Broin, 2001; several previously unpublished specimens that add Joyce et al., 2011; Pérez-García, 2012; Bardet et to the knowledge about both the shell anatomy and al., 2014). However, the British record of Pleurodira variability of this species, are also analyzed here. is very limited, being restricted to only a few rela- Photographs and schematic drawings of the more tively poorly known and poorly represented forms recently discovered specimens are also provided. from the lower Eocene (Gaffney et al., 2006; The detailed study of all these specimens supports Pérez-García and Chapman, 2017). Several their interpretation as belonging to a single spe- events of dispersion of this group of Gondwanan cies, represented by both juvenile and adult indi- origin to continents that were part of Laurasia are viduals. A reconstruction of the complete shell of known, and more specifically to Europe, both this taxon is done. The new combination Palemys during the Mesozoic and the Cenozoic (Pérez- bowerbankii (Owen, 1842) is proposed, and a neo- García, 2016a, 2017). Two successful lineages of type is designated. Additionally, this coastal bothre- Pleurodira are identified in the Eocene levels of mydid turtle is incorporated, for the first time, in a Europe: Bothremydidae and Podocnemididae phylogenetic analysis. (Lapparent de Broin, 2001). The podocnemidids Institutional abbreviations. GSM, British Geolog- are the best represented, showing a relatively wide ical Survey (BGS), Keyworth, Nottinghamshire, diversity. This group is represented in Continental UK; NHMUK, Natural History Museum, London, Europe both by freshwater forms (i.e., several spe- UK. cies of Neochelys Bergounioux, 1954) and by the recently described coastal genus Eocenochelus MATERIALS AND METHODS Pérez-García, Lapparent de Broin and Murelaga 2017 (Pérez-García et al., 2017). Eocenochelus is The detailed study of all the specimens of known in the Belgian, French, Spanish and Italian Palemys bowerbankii identified in this manuscript, records, as well as a single specimen from the including those previously published (except its upper Ypresian (early Eocene) of Bracklesham Bay holotype, now lost) and other unpublished speci- (Sussex, England) (Pérez-García et al., 2017; mens, is performed here. In this way, the shell of Pérez-García and Chapman, 2017; Pérez-García this generally considered problematic taxon can be and Smith, 2017). This recently identified fossil, characterized in detail. corresponding to an indeterminate species, rep- In order to evaluate the systematic position of resents the only podocnemidid specimen currently Palemys bowerbankii, it was coded in the data known in the British record (Pérez-García and matrix proposed by Pérez-García (2016a), which Chapman, 2017). pursues the study of the phylogenetic relationships Several shells and partial shells of coastal among the representatives of Bothremydidae bothremydid turtles have been found in the lower (Appendix 1. The recently described European Eocene British record, since the 19th century, from Eocene bothremydid Tartaruscola teodorii Pérez- the London Clay of the Isle of Sheppey (see Owen, García (2016b) is also included, following the codi- 1842, 1858; Owen and Bell, 1849). They were fication proposed by Pérez-García (2016b). Thus, attributed to four different species, ‘Platemys’ bow- the data matrix used here is composed of 180 cha- erbankii Owen, 1842, ‘Emys laevis’ Bell in Owen racters and 31 taxa (Appendix 2). and Bell, 1849, ‘Emys delabechii’ Bell in Owen and This data matrix was analyzed using TNT 1.0 Bell, 1849, and ‘Emys conybearii’ Owen, 1858. Wil- (Goloboff et al., 2008) in order to find the most par- liams (1954) indicated that all of them could repre- simonious trees (MPTs). The outgroup of this anal- sent a single species. No detailed review of these ysis is Proganochelys quenstedti Baur, 1887. A specimens has been made after the study of Wil- heuristic tree search (Traditional search) was used, liams (1954). In fact, although drawings of some of with 1000 replications of Wagner trees (using ran- the classic specimens were included in several dom addition sequences) followed by tree bisection works (presented by Owen and Bell, 1849, and recognition (TBR) as a swapping algorithm, saving reproduced in subsequent papers, see Owen, 100 trees per replication. To test the robustness of 1849; Williams, 1954), no photographs of any of the phylogenetic hypotheses, Bremer support and 2 PALAEO-ELECTRONICA.ORG bootstrap frequencies (absolute frequencies based (Podocnemis bowerbanki (Owen, 1842)) in Rayner on 1000 replications) values were also obtained et al. (2009). using TNT. Equal weighting was used for the analy- Emended diagnosis. Member of Bothremydini sis. characterized by the following combination of char- acters: relatively large shell, exceeding 45 cm; SYSTEMATIC PALEONTOLOGY elongated and elliptical carapace, its length being almost 1.4 times that of its width; slightly wider than Order TESTUDINES Batsch, 1788 long nuchal plate; seven neurals, the first five being Suborder PLEURODIRA Cope, 1864 notably longer than wide, especially the first, which Hyperfamily PELOMEDUSOIDES Cope, 1868 is more than twice as long as wide; antero-lateral Superfamily PODOCNEMIDOIDEA Cope, 1868 margins of the first vertebral scute reaching the Family BOTHREMYDIDAE Baur, 1891 second pair of peripherals; first vertebral as wide Infrafamily BOTHREMYDODDA Gaffney, Tong and as the second; first to third vertebrals wider than Meylan, 2006 long; overlap of the first pair of marginals on more Tribe BOTHREMYDINI Gaffney, Tong and Meylan, than half of the nuchal anterolateral margins; outer 2006 surface of the carapace and plastral plates lacking Subtribe FOXEMYDINA Gaffney, Tong and a well-developed ornamental pattern; long plastral Meylan, 2006 bridge, longer than the plastral lobes; broad plas- Genus PALEMYS Gray, 1870 tral lobes (the anterior being more than twice as Palemys bowerbankii (Owen, 1842) new wide as it is long), with sub-straight lateral margins; combination trapezoidal anterior lobe; short and wide anal
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