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A NEW LATE JURASSIC TURTLE from SPAIN: PHYLOGENETIC IMPLICATIONS, TAPHONOMY and PALAEOECOLOGY by BEN J

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A NEW LATE JURASSIC TURTLE from SPAIN: PHYLOGENETIC IMPLICATIONS, TAPHONOMY and PALAEOECOLOGY by BEN J [Palaeontology, Vol. 54, Part 6, 2011, pp. 1393–1414] A NEW LATE JURASSIC TURTLE FROM SPAIN: PHYLOGENETIC IMPLICATIONS, TAPHONOMY AND PALAEOECOLOGY by BEN J. SLATER1, MATI´AS REOLID2, REMMERT SCHOUTEN3 and MICHAEL J. BENTON3 1School of Geography, Earth and Environmental Sciences, University of Birmingham, Edgbaston, Birmingham B15 2TT, UK; e-mail: [email protected] 2Departmento de Geologı´a, Universidad de Jae´n, Campus Las Lagunillas sn, 23071 Jae´n, Spain; e-mail: [email protected] 3School of Earth Sciences, University of Bristol, Wills Memorial Building, Queen’s Road, Bristol BS8 1RJ, UK; e-mails: [email protected], [email protected] Typescript received 26 October 2010; accepted in revised form 27 May 2011 Abstract: The Jurassic was an important period in the evo- of the new taxon is hard to resolve, and it might be either a lution of Testudinata and encompasses the origin of many paracryptodire or a basal testudine, but it is distinct from clades, and this is especially true of Jurassic turtles from Wes- Plesiochelys. A complex taphonomic history is shown by a tern Europe. A new genus and species of Late Jurassic turtle, range of overlying grazing traces and bioerosion on the cara- Hispaniachelys prebetica gen. et sp. nov. from the upper Ox- pace. The carapace was subsequently overturned and buried fordian of the Prebetic (Southern Spain), is described on the ventrally up, terminating grazing activity, and was then bored basis of postcranial material. The specimen is the only known by sponges before final burial. Scanning electron microscopy tetrapod from the Mesozoic of the Prebetic and the oldest reveals phosphatic microspheroids associated with bacterial turtle from southern Europe. A mosaic of characters indicates decay in the vascular cavities of the cancellous bone, suggest- this is a new genus: it displays basal features including dorsal ing the carapace may have acted as a closed microenviron- epiplastral processes ⁄ reduced cleithra, no medial contact of ment in which decay-derived authigenic minerals formed. the extragulars and a long first thoracic rib, alongside derived characters including an absence of mesoplastra and the verte- Key words: Testudinata, Testudines, taphonomy, marine bral 3 ⁄ 4 sulcus crossing neural 5. The phylogenetic position turtles, Oxfordian, Spain. T he stratigraphic range of turtles (i.e. Testudinata sensu The earliest known turtle is Odontochelys semitestacea Joyce et al., 2004) extends from the Triassic to the present from the Late Triassic of Guizhou, China, which is day, with extant species occupying an array of niches, 220 myr old (Li et al. 2008), some 10 myr older than Pro- from terrestrial and aquatic to fully marine (Ernst and ganochelys quenstedti from Germany, previously regarded Barbour 1989). The evolution and development of the as the basal sister group to all other turtles (Gaffney and highly derived body plan of turtles has been a focus of Meylan 1988; Gaffney 1990; Joyce 2007). Modern turtles much attention (deBraga and Rieppel 1997; Lee 1997; are divided into two clades, Pleurodira and Cryptodira, Rieppel and Reisz 1999; Cebra-Thomas et al. 2005; Joyce based on the manner in which they withdraw their head et al. 2009; Rieppel 2009), which reflects the extreme and neck into the shell: pleurodires retract the head by modification of the basic tetrapod body plan in turtles, making a horizontal ‘S’ bend in the neck, whereas cryp- and their long-lasting success. New information has come todires make a vertical bend. These neck-folding modes from significant discoveries of early fossil stem turtles (Li are hard to determine in early fossil turtles, and the et al. 2008; Joyce et al. 2009) and study of the growth, assignment of some Jurassic taxa has been debated. For embryology and ontogeny of modern turtles (Burke 1989; example, Kayentachelys aprix from the Early Jurassic Hirayama 1998; Joyce and Gauthier 2004; Scheyer et al. (190 Ma) was previously thought to be the earliest cryp- 2008; Snover and Rhodin 2008; Rieppel 2009). Neverthe- todire (Gaffney et al. 1987), but more recent studies less, much is still not understood about early turtle evolu- (Joyce 2007) have suggested it is a more basal turtle and tion and divergence, particularly their relationship to that Pancryptodira, the clade including modern cryptod- other groups of amniotes (Laurin and Reisz 1995; Burke ires and their stem taxa, did not diverge until the Middle et al. 1996; Lee 1997; Zardoya and Meyer 2001). Jurassic (Danilov and Parham 2006, 2008; Sterli 2008). ª The Palaeontological Association doi: 10.1111/j.1475-4983.2011.01100.x 1393 1394 PALAEONTOLOGY, VOLUME 54 Complete turtle specimens are rare in the Jurassic cates that the turtle occurs in the Bimammatum Biozone (Joyce 2000), being confined usually to fragmentary (Olo´riz et al. 1999). The sediments were deposited in a remains of either the skull or shell in isolation. Much pre- mid-shelf environment on a carbonate epicontinental vious work on turtles has therefore focused on identifying shelf system. The marl-limestone rhythmites represent phylogenetic relationships through cranial (e.g. Gaffney fluctuations in carbonate productivity and delivery of si- 1975a, 1984) or shell characters (Joyce 2003). The skull liciclastic material derived from the proximal Iberian pal- displays wide disparity between species and contains aeomargin, which is defined by the extent of Iberian many diagnostic characters (Gaffney 1979), but it is often Massif tabular sediments immediately to the north (Olo´- not preserved, or only in isolation from carapace and riz et al. 2003, 2006; Reolid et al. 2010). The RGCH sec- plastron elements. Joyce (2007) provided a comprehensive tion has been heavily studied for its rich invertebrate character matrix of both cranial and postcranial charac- fauna (Olo´riz et al. 2002, 2003, 2006; Reolid 2003, 2008; ters, helpful here in that our specimen consists primarily Reolid et al. 2008), but vertebrates have not so far been of dermal elements of the shell, with some isolated verte- reported. brae. Late Jurassic turtles are distributed widely across Eur- ope, with specimens from Britain, Germany, France and METHODS Switzerland (Lapparent de Broin 2001). The majority of these have been assigned to the families Plesiochelyidae, The specimen was prepared by Remmert Schouten (RS) Eurysternidae and Thalassemydidae, but the distinctive- using both mechanical and chemical means. Pneumatic ness of these families has been questioned in recent air pens were used to remove the extensive limestone phylogenetic analyses (Joyce 2007). Late Jurassic turtles matrix, and the remaining material was dissolved in from the Iberian microcontinent include those from the 5 per cent acetic acid, buffered with calcium phosphate Lourinha˜ Formation (Kimmedigian–Tithonian) of wes- to protect the bone surface. In addition, a solution of tern Portugal (Antunes et al. 1988) and trackways from Paraloidª followed by a paraffin wax coating was the Late Jurassic Vega, Teren˜es, and Lastres formations painted onto the exposed areas of the bone prior to (Kimmeridgian) of Asturias, Spain (Avanzini et al. each acid treatment to further protect the bone from 2005). Turtles are poorly represented in the Oxfordian acid attack. Thin sections of the carapace were studied (Bardet 1995), with only an incomplete Plesiochelys and photographed under a plain light microscope to specimen from the Oxfordian of Germany (Kuhn assess the bone histology. 1949). Many fossil turtles are known from the Oxford For scanning electron microscopy (SEM), fragments of Clay of England, but this is largely Callovian in age carapace and indeterminate fossil bones were mounted (Martill and Hudson 1991). Outside Europe the Oxfor- and coated with gold and carbon. Secondary electron dian turtle Caribemys oxfordiensis has been found in images were made to study internal ultrastructure and Cuba (De la Fuente and Iturralde-Vinent 2001). Con- crystal morphology, and back-scattered electron (BSE) sidering their paucity in the Oxfordian, the discovery of imaging and energy-dispersive X-ray (EDX) analyses to a new turtle of this age from Spain is significant. obtain textural and chemical data. These analyses were The aim of this paper is to describe the new specimen performed using the FEI Quanta 400 at the Centro And- in detail, to identify it and determine its relationships, to aluz de Medio Ambiente (CEAMA, Granada). present information on its bone histology and taphonomy and to discuss its palaeoecology. SYSTEMATIC PALAEONTOLOGY GEOGRAPHICAL AND GEOLOGICAL Order TESTUDINATA Klein, 1760 (sensu Joyce et al. 2004) SETTING Infraorder TESTUDINES Batsch, 1788 The specimen was found by Matı´as Reolid (MR) in bed Hispaniachelys prebetica gen. et sp. nov. 62 (upper Oxfordian) of the Riogazas-Chorro (RGCH) Text-figure 2 section of the Lorente Formation of the External Prebetic, the northernmost part of the Betic Cordillera in southeast Derivation of name. Genus after the Latin for Spain and Greek Spain (Text-fig. 1). Bed 62 is within a 22-m sequence of for turtle, species after the Prebetic of Andalucı´a (Spain), where well-stratified limestone and marl rhythmites dominated the specimen was found. by limestones, which make up the Middle Oxfordian to Lower Kimmeridgian age Lorente Formation (Olo´riz Holotype. RGCHSP-62-52 (Text-fig. 2), housed in the Museo de ´ et al. 2002, 2003). The ammonite biostratigraphy indi- Paleontologıa of the Universidad de Granada, Spain. SLATER ET AL.:NEWJURASSICTURTLEFROMSPAIN 1395 TEXT-FIG. 1. Geological map of the Prebetic outcrops (Betic Cordillera) in the Sierra de Cazorla area (A) with location of the Riogazas-Chorro (RGCH) section (co-ordinates 3;0¢25¢¢– 35;52¢55¢¢ sheet 928 (Cazorla) of the National Topographic Map; Scale 1:50 000), and stratigraphic section (B) with the ammonite biostratigraphy for the Bimammatum Biozone within the Lorente Formation (based on Olo´riz et al. 1999); note black arrow indicates the level with the fossil turtle remains. Type stratum and locality. Bed 62 of the Lorente Formation; of preservation states.
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