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Reinterpretation of the Spanish Late “Hispaniachelys prebetica” as an indeterminate plesiochelyid

ADÁN PÉREZ-GARCÍA

Pérez-García, A. 2014. Reinterpretation of the Spanish “Hispaniachelys prebetica” as an indeterminate plesiochelyid turtle. Acta Palaeontologica Polonica 59 (4): 879–885.

A partial postcranial skeleton (carapace, plastron, and other poorly preserved elements) of a turtle, from the late Ox- fordian of the Betic Range of Spain, has recently been assigned to a new , Hispaniachelys prebetica. This is one of the few European turtle taxa reported from pre- levels, and the oldest turtle so far known from southern Europe. The character combination identified in that taxon (including the presence of cleithra, and single cervical scale) did not allow its assignment to , a group of very abundant and diverse in the Late Jurassic of Eu- rope. The revision of the single specimen assigned to this taxon led to the reinterpretation of some of its elements, being reassigned to Plesiochelyidae. This study confirms the presence of Plesiochelyidae in the Oxfordian. However, because the Spanish taxon does not present a unique combination of characters, it is proposed as a nomen dubium.

Key words: Testudines, Plesiochelyidae, Hispaniachelys prebetica, Oxfordian, Jurassic, Spain.

Adán Pérez-García [[email protected]], Centro de Geologia, Faculdade de Ciências da Universidade de Lisboa (FCUL), Edificio C6, Campo Grande, 1749-016 Lisbon, Portugal; Grupo de Biología Evolutiva, Facultad de Ciencias, UNED, C/ Senda del Rey, 9, 28040 Madrid, Spain.

Received 9 October 2012, accepted 29 May 2013, available online 5 June 2013.

Copyright © 2014 A. Pérez-García. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

lows to reconsider the and phylogenetic position of Introduction “Hispaniachelys prebetica”. It is proposed that it is a nomen dubium, the holotype being reassigned to Plesiochelyidae. The discovery of postcranial elements of a turtle from the Upper Oxfordian of the Prebetic area of the Betic Range (Southern Spain) has been recently reported (Slater et al. 2009, 2011). This finding is of particular interest since the Eu- Review of the anatomy and ropean pre-Kimmeridgian material of turtles is scarce (Lap- parent de Broin 2001; Anquetin and Claude 2008; Anquetin diagnosis of “Hispaniachelys et al. 2008). In addition, that specimen (RGCHSP-62-52) is prebetica” the oldest turtle so far known from southern Europe (Slater et al. 2011). A new and , Hispaniachelys prebetica The new study of RGCHSP-62-52 (Fig. 1), holotype and Slater, Reolid, Schouten, and Benton, 2011 was erected on the only specimen assigned to “Hispaniachelys prebetica”, al- basis of this material, diagnosed by a unique combination of lows reinterpretation of some of the characters used to di- characters. Its phylogenetic position was considered hard to agnose this taxon. This specimen preserves an epiplastron, resolve, being interpreted as either a paracryptodire or a basal on the right (Fig. 1C–E). This plate has not been described testudine, a priori excluding its assignment to Plesiochely- so far. The rear edge of this plate is subperpendicular to the idae, a very abundant in the Late Jurassic of Europe. axial plane. This morphology contrasts with the interpreta- The study of some elements that had not been previously tion proposed by Slater et al. (2011: fig. 3B), corresponding analysed (e.g., the right epiplastron); the reinterpretation of to the reconstruction of the ventral view of the plastron. The some characters, and new insights into the morphology of visceral view of this element (Fig. 1D, E) shows that, con- some elements hitherto poorly described in the traditional trary to the previous interpretation, this taxon lacked dorsal basal members of Eucryptodira from the European record al- epiplastral processes (cleithra) and instead the structure in

Acta Palaeontol. Pol. 59 (4): 879–885, 2014 http://dx.doi.org/10.4202/app.2012.0115 880 ACTA PALAEONTOLOGICA POLONICA 59 (4), 2014 question is probably a result of the aberrant morphology of Phylogenetic position of the lateral margin of the right hyoplastron. The presence of this element motivated Slater et al. (2011) to exclude assign- “Hispaniachelys prebetica” ment of RGCHSP-62-52 to Plesiochelyidae, a family that traditionally includes several genera from the Late Jurassic Slater et al. (2011) explored the systematic position of RGC- of Europe: Craspedochelys Rütimeyer, 1873, HSP-62-52 using a modified version of the data matrix of Rütimeyer, 1873, Portlandemys Gaffney, 1975a, Tholemys Joyce (2007) as proposed by Anquetin et al. (2008). The Andrews, 1921, and Tropidemys Rütimeyer, 1873 (Lappar- revision of the specimen RGCHSP-62-52 performed here ent de Broin 2001; Lapparent de Broin et al. 1996). allows emendation of some of the character coding (see Ap- As indicated by Slater et al. (2011), the nuchal plate is pendix 1). Because RGCHSP-62-52 corresponds to a partial almost complete but poorly preserved as its dorsal surface is postcranial skeleton, it is coded for only 43 of the 139 charac- heavily weathered and obscured by erosional pits, especially ters of this matrix. This new phylogenetic analysis has been in its anterior half (Fig. 1B). Therefore, the hypothesis pro- conducted using TNT v. 1.0 (Goloboff et al. 2008), the hy- posed by Slater et al. (2011) about the presence of a single pothetical ancestor being the outgroup, and the three “rogue” cervical scale cannot be confirmed. It cannot be excluded taxa indicated by Joyce (2007) being excluded: Portlande- that this specimen shares with the members of Plesiochely- mys mcdowelli Gaffney, 1975a, Sandownia harrisi Meylan, idae the presence of three cervicals. Moody, Walther, and Chapman, 2000, and Mongolemys ele- The review of the plastral elements of RGCHSP-62-52 gans Khozatsky and Młynarski, 1971. This analysis has been reveals new information regarding the morphology of sever- performed using a traditional search, in which, as Slater et al. (2011) did, a tree-bisection algorithm, with 5000 replicates, al scales (Fig. 1C2). Although the presence of short pectoral scales was considered in the diagnosis of “Hispaniachelys was applied. All characters were considered unordered. prebetica”, the preserved elements of the plastron show that The phylogenetic analysis resulted in 37,501 most parsi- medially, this pair of scales was probably longer than the pair monious trees of 370 steps (consistency index = 0.466; reten- of abdominals (Fig. 1C). The presence of a partially sinusoi- tion index = 0.811; rescaled consistency index = 0.378) (Fig. dal humero-pectoral sulcus is refuted; it is subperpendicular 2). The resulting strict consensus topology basically coincides to the axial plane, slightly convex medially. with that proposed by Slater et al. (2011), RGCHSP-62-52 be- Other characters present in RGCHSP-62-52, which were ing obtained in a polytomy composed of 30 branches, corre- not part of the diagnosis of “Hispaniachelys prebetica”, but sponding to some taxa identified in other papers as members which had been described and incorporated in the reconstruc- of the stem group of Testudines and the representatives of the tion of that taxon (Slater et al. 2011: fig. 3), are also reinterpret- crown group (see Joyce 2007; Anquetin 2008, 2012; Sterli ed here. As indicated by Slater et al. (2011), the internal mor- and de la Fuente 2011; Pérez-García et al. 2014). However, phology of the bridge peripherals is often not described and although the majority-rule tree obtained by Slater et al. (2011) figured in extinct turtles. This lack of information generated an placed RGCHSP-62-52 as a member of , asso- erroneous interpretation of the arrangement of the disarticulat- ciated with the baenid clade, the majority-rule tree obtained ed bridge peripherals of RGCHSP-62-52. Thus, the groove on here does not include RGCHSP-62-52 between the repre- the contact of the peripherals and the distal region of the dorsal sentatives of Paracryptodira, from which it differs, among ribs was provisionally interpreted as the sutural surface of the other characters, by the absence of mesoplastra (this character carapace and plastron. Therefore the dorsal region of these being one of the characters that diagnosed the node Paracryp- peripheral plates was interpreted as the ventral region (Slater todira in the majority-rule tree, and also the nodes et al. 2011: fig. 4). This also led to an erroneous interpretation and in the strict consensus tree). In this tree, of the position of these elements in the peripheral ring: the RGCHSP-62-52 is located in a basal polytomy of Testudines, plate identified as the sixth right peripheral (Slater et al. 2011: where the other members of Plesiochelyidae included in this fig. 4F, H, L) is interpreted here as the seventh right peripheral analysis are also situated (“Thalassemys” moseri Bräm, 1965 (Fig. 1I–M) and has a fully consistent morphology with the and Plesiochelys solodurensis Rütimeyer, 1873). corresponding element on the left side (Fig. 1F–H). Similarly, RGCHSP-62-52 does not have autapomorphies in this the fifth or sixth right peripheral (Fig. 1N–P) was identified as analysis. In fact, the 43 characters coded for this taxon are the fourth right peripheral (Slater et al. 2011: fig. 4E, G, K). compatible with those coded for the two members of Plesio- Slater et al. (2011) interpreted a unique morphology of the chelyidae included in this analysis (both taxa including not inframarginal scales due to the problems in the orientation of only information on the postcranial skeleton, but also on the these plates (Slater et al. 2011: figs. 3B, 4K, L). However, the ). limited information available on these scales (Fig. 1C, G, H, J, The poor resolution of phylogenetic relationships among K, N) is compatible with that of members of Plesiochelyidae, the members of the stem group of , and between which possess a complete series of inframarginal scales, all Pancryptodira and Panpleurodira, offers an incomplete reso- of them with a similar width, situated from the axillary to the lution of several of the major clades (see Joyce 2007; Anque- inguinal notch, and therefore completely blocking any contact tin 2008, 2012; Sterli and de la Fuente 2011). In this sense, between the carapacial and plastral scales (Joyce 2007). the phylogenetic relationships among the representatives of PÉREZ-GARCÍA—JURASSIC TURTLE HISPANIACHELYS PREBETICA REVISITED 881

A1 B1 C1 D E

F1

G1

H1 5cm

peripheral 1 extragular A 2 costal 1 B2 C2 marginal 3 nuchal humeral pleural 1 epiplastron marginal 7 peripheral 4 marginal 1 pleural 2 pectoral pleural 3 vertebral 1 costal 3 neural 1 F2 neural 4 hyoplastron vertebral 3 marginal 8 G 2 abdominal

hypoplastron peripheral 7 pleural 3 H marginal 9 inframarginal 2 neural 7 femoral marginal 8 suprapygal 1 xiphiplastron marginal 7 pleural 10 inframarginal costal 8 suprapygal 2 anal Rs Rs I1 J1 K1 L N1 O

pleural 3 pleural 2 marginal 8 I 2 J2 K 2 M N2 P marginal 7 marginal 8 marginal 7

Fig. 1. RGCHSP-62-52, shell of the holotype of “Hispaniachelys prebetica” from late Oxfordian of the Betic Range of Spain, interpreted as plesiochelyid turtle herein. Carapace in dorsal view (A); most-anterior region of the carapace in dorsal view (B); plastron in ventral view (C); right epiplastron in dorsal (D) and posterior (E) views; seventh left peripheral in dorso-lateral (F), latero-ventral (G), and ventral (H) views; seventh right peripheral in dorso-lateral (I), latero-ventral (J), ventral (K), medial (L), and posterior (M) views; fifth or sixth right peripheral in latero-ventral (N), medial (O), and anterior (P) views. Photographs (A1–C1, F1–K1, N1) and explanatory drawings (A2–C2, F2–K2, N2). Rs, groove generated by the distal region of the dorsal ribs on the visceral surface of the peripherals. 882 ACTA PALAEONTOLOGICA POLONICA 59 (4), 2014

hypothetical ancestor hypothetical ancestor quenstedti Proganochelys quenstedti Proterochersis robusta Proterochersis robusta Palaeochersis talampayensis Palaeochersis talampayensis africanus Australochelys africanus aprix Kayentachelys aprix Condorchelys antiqua Condorchelys antiqua 80 Eileanchelys waldmani Eileanchelys waldmani Heckerochelys romani Heckerochelys romani 77 platyceps 51 Meiolania platyceps Mongolochelys efremovi Mongolochelys efremovi bajazidi 84 Kallokibotion bajazidi RGCHSP-62-52 Testudinata RGCHSP-62-52 Plesiochelys solodurensis Plesiochelys solodurensis “Thalassemys” moseri “Thalassemys” moseri 96 Dorsetochelys delairi Dorsetochelys delairi bullockii Pleurosternon bullockii plicatulus Glyptops plicatulus 70 Pleurosternidae Dinochelys whitei Pleurosternidae Dinochelys whitei Paracryptodira eximius 79 Neurankylus eximius 94 Trinitichelys hiatti Trinitichelys hiatti Testudines Baenidae antiqua Plesiobaena antiqua Boremys pulchra Boremys pulchra arenosa Baena arenosa Chisternon undatum Baenidae Chisternon undatum 59 Solnhofia parsonsi Solnhofia parsonsi gaffneyi Santanachelys gaffneyi Xinjiangchelys junggarensis Xinjiangchelys junggarensis 58 Mesodermochelys undulatus Mesodermochelys undulatus latiremis Toxochelys latiremis Panchelonioidea 75 58 75 Dermochelys coriacea Dermochelys coriacea Caretta caretta 60 Caretta caretta Chelonia mydas Chelonia mydas Panchelonioidea Kirgizemys hoburensis Kirgizemys hoburensis 58 77 Judithemys sukhanovi Judithemys sukhanovi Cryptodira 57 Dracochelys bicuspis Dracochelys bicuspis 97 lens Sinemys lens leios Ordosemys leios Platysternon megacephalum Platysternon megacephalum 58 81 Protochelydra zangerli Protochelydra zangerli Macroclemys temminckii 63 Macroclemys temminckii Panchelydridae serpentina Chelydra serpentina polyphemus Gopherus polyphemus Geoclemys hamiltonii Geoclemys hamiltonii 58 58 Chrysemys picta Chrysemys picta dentata Elseya dentata 58 66 oblonga 50 Chelodina oblonga geoffroanus Phrynops geoffroanus Notoemys laticentralis Notoemys laticentralis 64 58 Platychelys oberndorferi Platychelys oberndorferi 58 Caribemys oxfordiensis Caribemys oxfordiensis Pelomedusa subrufa Pelomedusa subrufa 58 58 80 Panpleurodira 58 Erymnochelys madagascariensis Erymnochelys madagascariensis expansa Panpleurodira Podocnemis expansa Emarginachelys cretacea Emarginachelys cretacea 65 Baptemys wyomingensis Baptemys wyomingensis Dermatemys mawii Dermatemys mawii 58 Hoplochelys crassa Hoplochelys crassa triporcatus 86 Staurotypus triporcatus 65 odoratus 96 Trionychoidea flavescens Kinosternon flavescens Kinosternoidea beatus Adocus beatus Zangerlia neimongolensis 72 Zangerlia neimongolensis variolosa Basilemys variolosa Peltochelys duchastelii Peltochelys duchastelii ornata Anosteira ornata Carettochelys insculpa 87 Carettochelys insculpa spinifera Trionychia 99 Apalone spinifera A punctata B Lissemys punctata Fig. 2. Hypothesized phylogenetic position of the specimen RGCHSP-62-52, holotype of “Hispaniachelys prebetica” from late Oxfordian of the Betic Range of Spain, based on the modified data set of Joyce (2007) as proposed by Anquetin et al. (2008). A. Majority rule tree. Values are percentages under 100% obtained in this analysis. B. Strict consensus tree with bootstrap values (1000 replicates, only values over 50% shown). Plesiochelyidae, and between them and other nodes, are cur- TNT v. 1.0, using a traditional search in which a tree-bisection rently poorly understood. This was, in part, due to the limited algorithm, with 5.000 replicates, is applied. All characters are information about the members of Plesiochelyidae so far considered unordered and equally weighted. Some previously considered in global phylogenetic analyses. not processed characters are coded for the taxon Hylaeochelys There are other global analyses of turtle relationships Lydekker, 1889 in the first of these two analyses: character 67 where information about Plesiochelyidae is combined (inframarginals) as 0 (four or three pairs present), character 70 into a single terminal taxon, called Plesiochelyidae. RGC- (epiplastron) as 0 (moderately broad), character 71 (epiplas- HSP-62-52 is included here in two of them, those proposed tra subdivided by entoplastron) as 1 (no). In addition, the by Hirayama et al. (2000) and by Gaffney et al. (2007) (see character 68 (femoral-anal sulcus reaching hypoplastron) of Appendix 1). These two analyses were also processed with Plesiochelyidae is recoded as 0 and 1 (no and yes). Character PÉREZ-GARCÍA—JURASSIC TURTLE HISPANIACHELYS PREBETICA REVISITED 883 109 (pectoral scale) of Plesiochelyidae is coded as 0 (not on Proganochelys epiplastron), and character 110 (femoral-anal sulcus) of the Australochelyidae same taxon is recoded as 0 and 1 (not reaching hypoplastron 100 and reaching hypoplastron) in the second analysis. Both the Proterochersis Kayentachelys coding of the 16 characters of RGCHSP-62-52 processed in 63 the analysis of Hirayama et al. (2000) (data matrix consist- Otwayemys Sinochelyidae ing of 76 characters), and the 21 characters processed in the Mongolochelys analysis of Gaffney et al. (2007) (data matrix consisting of A 111 characters) coincide with the states of those characters Tretosternon present in Plesiochelyidae, differing from the coding of the Kallokibotion members of other nodes. The strict consensus tree obtained Paracryptodira from the first of these analyses generates a polytomy in which RGCHSP-62-52 RGCHSP-62-52, Plesiochelyidae, Hylaeochelys, and a node Hylaeochelys which includes the remaining members of Eucryptodira (sen- Plesiochelyidae Brodiechelys su Joyce et al. 2004) are located (Fig. 3A). These taxa are 73 grouped by several apomorphies, one of them observed in Sinemydidae the preserved material of RGCHSP-62-52: the absence of 50 Chelydroidea mesoplastra (character 74). The coding of the characters of 56 RGCHSP-62-52 differs from that of Hylaeochelys due to the Chelonioidea absence of vertebral scales much broader than costal scales Trionychoidea (character 59) and the presence of a femoro-anal sulcus reach- ing the hypoplastron (character 68). The node that groups the Sinapsida/Diapsida Proganochelys other representatives of Eucryptodira is diagnosed by sev- 100 eral character states, only one of them corresponding to the Australochelys elements preserved in RGCHSP-62-52: plastral buttresses 73 Palaeochersis Megapleurodira reaching only peripherals (character 66). In the second analy- 64 62 sis, RGCHSP-62-52, Plesiochelyidae and Solnhofia Gaffney, B Proterochersis 1975b (a member of Eurysternidae, another group of basal 72 Kayentachelys eucryptodires, also exclusive of the European Upper Jurassic Mongolochelys record) are grouped in a node diagnosed by several charac- Kallokibotion ters corresponding to elements not preserved in the specimen Baenidae analysed here (Fig. 3A). The node grouping RGCHSP-62-52 Pleurosternidae and Plesiochelyidae is diagnosed by the presence of sutured Xinjiangchelys carapace-plastron attachment (character 96) and plastral but- Solnhofia tresses reaching the costals (character 97). RGCHSP-62-52 Therefore, all the relatively limited character states of Plesiochelyidae RGCHSP-62-52 codified in all the analyses are the same Otwayemys as those encoded to both Plesiochelyidae (in the analyses in Chubutemys Niolamia which Plesiochelyidae is encoded as a single taxon) and all 67 the coded representatives of Plesiochelyidae: Plesiochelys 76 Ninjemys solodurensis and “Thalassemys” moseri. In fact, the charac- Meiolania ters available in RGCHSP-62-52 (both those considered in Ordosemys the cladistic studies and those not included in these global Sinemys analyses but incorporated in other more traditional system- Dracochelys atic analyses) indicate several character states that constitute Judithemys a unique combination of the eucryptodire group Plesioche- Hangaiemys lyidae (sensu Lapparent de Broin et al. 1996): nuchal notch 74 Testudinoidea very small or absent; sub-trapezoidal morphology of the nu- Trionychoidea chal, in which the most-anterior region of their lateral mar- Chelonioidea gins generally changes its curvature, resulting in diverging margins; anterior peripherals shorter and wider than that of the members of Eurysternidae, with its anterior margin being Fig. 3. Strict consensus trees showing the hypothesized phylogenetic posi- tions of the specimen RGCHSP-62-52, holotype of “Hispaniachelys preb- more delayed than that of the nuchal; presence of at least two etica”, based on the data set of: A, Hirayama et al. (2000) (trees retained: suprapygal plates; hexagonal vertebrals, no much broader 2, tree length: 133, CI: 0.624, RI: 0.745); B, Gaffney et al. (2007) (trees than pleural scales; absence of overlapping of the marginal retained: 2, tree length: 225, CI: 0.569, RI: 0.744). Bootstrap values (1000 scales onto the costal plates; absence of fontanelles in the car- replicates) over 50% are shown. 884 ACTA PALAEONTOLOGICA POLONICA 59 (4), 2014 apace; more domed shell than in Eurysternidae and Thalas- supported by the characters described in the single specimen semydidae; sutured carapace-plastron attachment; buttresses attributed to it, it cannot be either validated or synonymized of the hyoplastra and hypoplastra in contact with the costal with other plesiochelyid representatives, this taxon is here series; absence of bridge fontanelles but presence of a central identified as a nomen dubium. Several characters observed plastral fontanelle in many inferred adult individuals. in RGCHSP-62-52, but not taken into account in the diag- nosis of “Hispaniachelys prebetica”, are shared with several members of Plesiochelys (e.g., the overall shape of its shell, Taxonomy of “Hispaniachelys the absence of a sagittal keel, the width of the vertebral scales). Due to the limited information on the diagnosed prebetica” characters of several genera and species of Plesiochelyidae, and to the scarce information available on RGCHSP-62-52, As indicated, the diagnosis of “Hispaniachelys prebetica” this specimen is considered here as an indeterminate mem- (sensu Slater et al. 2011) does not include autapomorphies. ber of Plesiochelyidae. Review of several members of this In fact, many taxa that could belong to Plesiochelyidae, de- clade is needed to establish the phylogenetic position of this termined by scarce material, and whose validity has not been specimen. assessed, have been described in the Late Jurassic of Europe The record of Plesiochelyidae, a group of coastal marine (e.g., Plesiochelys hannoverana [Maack, 1869], “” turtles, is abundant in the Kimmeridgian– levels, lindenensis Maack, 1869, “” dollfussi Lennier, 1870, having being recognized in several European countries (Lap- “Chelonides” robusta Portis, 1878, Plesiochelys normandi- parent de Broin 2001; Lapparent de Broin et al. 1996). In the cus Bergounioux, 1937). As previously indicated, the rein- Iberian Peninsula this clade has been identified both in the terpretation of some of these characters shows that RGC- Kimmeridgian of Spain and Portugal (Antunes et al. 1988; HSP-62-52 lacked epiplastral process, and the length of the pectoral scales and the morphology of the humero-pectoral Pérez-García et al. 2010), and in the Tithonian of both coun- groove were not different to those present in the members of tries (Pérez-García et al. 2008; Pérez-García et al. 2013). The Plesiochelyidae. Due to its preservation, the number of cervi- currently known Iberian record includes representatives of cal scales is not known. The other characters present in RGC- Craspedochelys, Plesiochelys, and Tropidemys (Antunes et HSP-62-52 and included in the diagnosis of “Hispaniachelys al. 1988; Lapparent de Broin 2001; Pérez-García et al. 2008; prebetica” (see Slater et al. 2011) are shared with all the taxa Pérez-García et al. 2013). The abundance, diversity and rel- traditionally attributed to Plesiochelyidae for which the shell ative wide distribution of this European clade during the is known (e.g., Craspedochelys, Plesiochelys, Tholemys, Kimmeridgian and Tithonian (Britain, France, Switzerland, Tropidemys): the moderately large size of the shell, lacking , Portugal, Spain) means that the discovery of Ple- fontanelles between the costal and peripheral plates; the mor- siochelyidae in the upper Oxfordian of the Iberian Peninsula phology of the neural plates (not all being hexagonal, but the was expected. first one sub-rectangular and the others hexagonal, except the last one, whose morphology is irregular); the presence of eight neurals (number generally subject to intraspecific Acknowledgements variability in basal eucryptodires, as noted, for example, by Lapparent de Broin et al. 1996 and Pérez-García 2012); the The author thanks Francisco Javier Rodríguez Tovar (University of number of costals (not eight plates but eight pairs of plates); Granada, Spain) and the authors of the first paper on RGCHSP-62-52, the sub-trapezoidal morphology of the nuchal (in which the especially Matías Reolid (University of Jaen, Spain), for access to the cited specimen, as well as to other documentation and information most-anterior region of their lateral margins changes its cur- related to it; and Mike Benton (University of Bristol, UK), Jean-Paul vature, resulting in diverging margins); the diamond shape of Billon-Bruyat (Office de la Culture, République et Canton du Jura, the entoplastron that does not separate the anterior portions Porrentruy, Switzerland), Walter Joyce (University of Tübingen, Ger- of the epiplastra; the arrangement of the sulcus generated be- many), Márton Rabi (University of Tübingen, Germany), and Juliana tween the third and fourth vertebrals crossing the fifth neural; Sterli (Museo Paleontológico Egidio Feruglio, Trelew, Argentina) for and the presence of anal scales confined to the xiphiplastra comments and suggestions. The Willi Hennig Society sponsors TNT (actually in contact with the hypoplastra in RGCHSP-62-52). software. This last character may be subject to individual variability, in which these scales in different specimens are limited to the xiphiplastra, in contact with the most posterior region References of the hypoplastra, or overlapping onto the hypoplastra (see examples of the three situations in Craspedochelys jaccardi Andrews, C.A. 1921. On a new chelonian from the Kimmeridge Clay of (Pictet, 1860: pls. 1, 4.4, 3, respectively; Lapparent de Broin Swindon. Annals and Magazine of Natural History 7: 145–153. Anquetin, J. 2011. Reassessment of the phylogenetic interrelationships of et al. 1996), and of the first and last in Plesiochelys solodu- basal turtles (Testudinata). Journal of Systematic Palaeontology 10: rensis (Bräm 1965: pls. 5.2, 5.4, respectively). Since the 3–45. taxonomic validity of “Hispaniachelys prebetica” cannot be Anquetin, J. and Claude, J. 2008. 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Appendix 1

Scored characters for the specimen RGCHSP-62-52 in the data matrices used here.

Data matrix of Joyce (2007) as proposed by Anquetin et al. (2008): ?????????? ?????????? ?????????? ?????????? ?????????? ?????????0 010010000? 2101010111 0001210000 0001000000 ?????????? ?????0???2 ?????????? ????????0 Data matrix of Hirayama et al. (2000): ?????????? ?????????? ?????????? ?????????? ?????????? ??????1010 ?01?010110 1?110? Data matrix of Gaffney et al. (2007): ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ???00??0?? ?????????? ???????00? ?1?2101120 0010??0001?