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A New Suoid with Tubulidentate, Hypselorhizic Cheek Teeth from the Early Miocene of Córcoles, Spain

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A New Suoid with Tubulidentate, Hypselorhizic Cheek Teeth from the Early Miocene of Córcoles, Spain SPANISH JOURNAL OF PALAEONTOLOGY A new suoid with tubulidentate, hypselorhizic cheek teeth from the early Miocene of Córcoles, Spain Martin PICKFORD1* & Jorge MORALES2 1 CR2P/UMR 7207 – Sorbonne Université – MNHN, CNRS, UPMC, Université Paris VI, T.46-56, E.5, case 104, 4 Place Jussieu, 75252 Paris cedex 05, France; [email protected] 2 Departamento de Paleobiología, Museo Nacional de Ciencias Naturales-CSIC – C/José Gutiérrez Abascal 2, 28006, Madrid, Spain; [email protected] * Corresponding author Pickford, M. & Morales, J. 2018. A new suoid with tubulidentate, hypselorhizic cheek teeth from the early Miocene of Córcoles, Spain. [Un nuevo suido con dientes yugales tubulidentados e hipselorrizos del Mioceno inferior de Córcoles, España]. Spanish Journal of Palaeontology, 33 (2), 321-344. Manuscript received 9 July 2018 © Sociedad Española de Paleontología ISSN 2255-0550 Manuscript accepted 17 October 2018 ABSTRACT RESUMEN Excavations at Córcoles, Guadalajara, Spain (MN 4) in Las excavaciones realizadas en el yacimiento de Córcoles, October 1987, yielded a mandible and several isolated teeth Guadalajara, España (MN 4) han suministrado una mandíbula of a strange suiform with tubulidentate cheek teeth, initially y algunos dientes aislados de un extraño suido con los interpreted to be the remains of Tubulidentata. The specimens dientes yugales tubulidentados, que inicialmente fueron are complete enough to remove any doubt about which interpretados como pertenecientes a un Tubulidentata Order (Artiodactyla) and Superfamily (Suoidea) they belong (cerdo hormiguero, Orycteropus). Los ejemplares son lo to. They are here attributed to the Family Doliochoeridae, sufi cientemente completos como para que no hayan dudas being related to the genera Bransatochoerus (MP 30) and de su pertenencia al Orden Artiodactyla, y dentro de él a Lorancahyus (MN 2) both of which have hypsorhizic cheek la Superfamilia Suoidea. En este trabajo son atribuidos a teeth, the latter with tubules in the teeth, but not to the la Familia Doliochoeridae, relacionándoles con los géneros fully-expressed extent seen in the fossils from Córcoles and Brasantochoerus (MP 30) y Lorancahyus (MN 2), ambos con the molars retain a distinct and large pulp cavity, which is dientes yugales hipsorrizos, éste último género con túbulos absent in the Córcoles species. It is inferred that these suoids dentarios, pero que no alcanzan un desarrollo tan extenso comprise an evolutionary lineage which went extinct in the como se aprecia en los fósiles de Córcoles, de manera que los early Miocene (ca MN 4b or MN 5) and that the dental molares aún retienen la cavidad pulpar, que está ausente en la specialisations imply ingestion of food to which soil was especie de Córcoles. Se infi ere que estos suoideos formaron adhering, which caused excessive abrasion of the teeth, not una línea evolutiva que se extinguió en el Mioceno inferior only occlusally, but also on the lingual aspects of the cheek (MN 4 o MN 5) y que la especialización dentaria implicaba teeth. Examination of the occlusal and lingual surfaces of the la ingesta de comida con particulas del suelo adheridas, lo teeth confi rm the presence of many pits and large scratches que causaba una abrasión intensa de los dientes, no solo en el https://doi.org/10.7203/sjp.33.2.13606 322 PICKFORD & MORALES on them, in contrast to the buccal and distal surfaces of the plano oclusal si no también en la muralla lingual de la serie teeth which have few scratches or none, respectively. dental. El examen de las superfi cies linguales y oclusales de los dientes confi rma la presencia de numerosos hoyos y Keywords: Hyposorhizy, hypselorhizy, tubulidenty, dental grandes rayaduras, en contraste con las superfi cies labiales y adaptations, Suoidea, early Miocene, Western Europe. distales de los dientes que tienen pocas o ninguna rayadura. Palabras clave: Hyposorrizo, hipselorrizo, tubulidentado, adaptaciones dentales, Suoidea, Mioceno inferior, Europa Occidental. 1. INTRODUCTION The Iberian Peninsula has a rich and extensive Cainozoic gomphotheres, which are among the earliest known from fossil record, and over the years it has yielded abundant Europe, implying a phase of dispersal of African large and varied fossils including many mammals. Its mid- mammals into Europe during the early Miocene (MN 4a). latitude position between Africa to the south and the rest In 1987, remains of an enigmatic medium-sized mammal of Europe to the north and east is of special signifi cance were found, characterised by its tubulidentate cheek tooth in that its fossil record preserves evidence of faunal structure. A mandible and several isolated cheek teeth were and fl oral dispersal between the continents (Pickford & collected and were interpreted by Alférez et al. (1988a, Morales, 1994). 1988b, 1999) to belong to Tubulidentata (Aardvarks, of The fossiliferous site of Córcoles, Guadalajara (Fig. 1), which the only surviving example is Orycteropus afer) was fi rst mentioned by Crusafont & Quintero (1970) but and the presence of this mammal in the Córcoles fauna extensive sampling only took place much later. In the was taken to provide additional evidence of mammalian 1980’s, excavations at Córcoles yielded a varied assortment dispersal from Africa into Europe during MN 4. The of vertebrates (Table 1) (Alférez & Brea, 1981; Alférez possibility that the Córcoles material belonged to Suoidea et al., 1981, 1982, 1988a, 1988b, 1999; Maldonado was raised by Pickford (1993) on the basis of some dento- & Alférez, 1990; Maldonado et al., 1983) including gnathic similarities that it shares with Lorancahyus, a genus fi rst described from older deposits in Spain (Loranca and Moheda, MN 2) (Figs 1-2). Figure 2. Biostratigraphic correlations of European localities that have yielded remains of Orycterochoerinae subfam. nov. Figure 1. Geographic distribution of Orycterochoerinae. The record from Navarrete del Rio is provisional. A NEW SUOID WITH TUBULIDENTATE, HYPSELORHIZIC CHEEK TEETH FROM THE EARLY MIOCENE OF CÓRCOLES, SPAIN 323 Table 1. Mammalian fauna associated with Orycterochoerus belong to Suoidea rather than to Tubulidentata. Pickford alferezi gen. et sp. nov. at Córcoles, Spain. (2011, 2016, 2017) described additional fossils of this lineage from St Gérand-le-Puy and Montaigu-le-Blin, Lagomorpha Lagopsis penai France (MN 2, originally attributed to Lorancahyus «Insectivora» Galerix sp. hypsorhizus by Pickford, 2016) and Pickford & Hugueney Amphechinus sp. (2018) described suoid fossils from Coderet, France Miosorex sp. (late Oligocene, MP 30 = MN 0) (Figs 1-2) which have Chiroptera Pseudorhinolophus sp. hypsorhizic teeth with enamel caps that rapidly wear away leaving only the roots as the part functioning during Genus indet. mastication. A peculiar aspect of the Coderet teeth, which Rodentia Megacricetodon primitivus are attributed to Bransatochoerus elaverensis (Viret, Democricetodon decipiens 1928a) is that they continue to erupt when heavily worn, Democricetodon sacedonensis thereby maintaining the occlusal surface of each tooth at Ligerimys sp. the same level as the other less-worn teeth in the cheek- Pseudodryomys ibericus tooth battery, unlike most mammals in which heavily Simplomys simplicidens worn molars decrease in height relative to the occlusal Simplomys meulenorum surface because the roots cease growing after maturity. Simplomys julii Bransatochoerus is known from skulls, mandibles and Peridryomys murinus substantial parts of the associated post-cranial skeleton (Pickford, 2016, 2017) and it is evident from the terminal Microdryomys legidensis phalanges that it was using its digits to excavate in the soil, Armantomys aragonensis perhaps to dig burrows or in search of underground food Armantomys jasperi resources (or both). The types and quantity of heavy wear Atlantoxerus cf blacki scars on the cheek teeth (pits, scratches, grooves) attest to Heteroxerus rubricati the mastication of soil particles with the food. Equidae Anchitherium corcolense The available fossil evidence indicates that a small Chalicotheriidae Phyllotillon naricus «peccary-like» suoid lineage that existed in Western Rhinocerotidae Hispanotherium matritense Europe during the Oligocene, developed hypsorhizic cheek Protaceratherium platyodon teeth with strongly fused roots (Bransatochoerus, MP Cainotheriidae Cainotherium miocaenicum 30) which eventually started developing tubular dentine rods within the fused roots (Lorancahyus, MN 2) and Suidae Eurolistriodon adelli which, by MN 4 times (Córcoles) (Fig. 2) culminated in Listriodon retamaensis a species with well-developed tubular dentine structure in Doliochoeridae Orycterochoerus alferezi gen. et sp. nov. the cheek teeth and precocious elimination of the enamel Ruminantia Dorcatherium crassum caps of the molars. In this lineage, during the life of the Procervulus dichotomus individuals, the bulk of mastication of food was achieved Hydropotopsis sp. by the roots rather than by the dental crowns which wore Triceromeryx pachecoi away at an early age, as was explained by Pickford & Eotragus artenensis Morales (1998) for Lorancahyus and by Pickford (2016) Proboscidea Gomphotherium angustidens for Bransatochoerus. The Córcoles suoid continued with this evolutionary trend to such an extent that its cheek Carnivora Pseudaelurus quadridentatus teeth superfi cially resemble those of aardvarks (Alférez et Amphicyon major al., 1999). Furthermore, the roots are ever-growing, here Plithocyon armagnacensis defi ned as hypselorhizic, in contrast to hypselodont
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