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Canidae, Carnivora, Mammalia) from Early Pliocene of Çalta, Turkey

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Canidae, Carnivora, Mammalia) from Early Pliocene of Çalta, Turkey geodiversitas 2019 ● 41 ● 18 né – Car ig ni e vo P r e e s n a o f h t t p h h é e e t S C C l l e e a a n i i r o o z o m i e c – M DIRECTEUR DE LA PUBLICATION : Bruno David, Président du Muséum national d’Histoire naturelle RÉDACTEUR EN CHEF / EDITOR-IN-CHIEF : Didier Merle ASSISTANTS DE RÉDACTION / ASSISTANT EDITORS : Emmanuel Côtez ([email protected]) ; Anne Mabille MISE EN PAGE / PAGE LAYOUT : Emmanuel Côtez COMITÉ SCIENTIFIQUE / SCIENTIFIC BOARD : Christine Argot (MNHN, Paris) Beatrix Azanza (Museo Nacional de Ciencias Naturales, Madrid) Raymond L. Bernor (Howard University, Washington DC) Alain Blieck (chercheur CNRS retraité, Haubourdin) Henning Blom (Uppsala University) Jean Broutin (UPMC, Paris) Gaël Clément (MNHN, Paris) Ted Daeschler (Academy of Natural Sciences, Philadelphie) Bruno David (MNHN, Paris) Gregory D. Edgecombe (The Natural History Museum, Londres) Ursula Göhlich (Natural History Museum Vienna) Jin Meng (American Museum of Natural History, New York) Brigitte Meyer-Berthaud (CIRAD, Montpellier) Zhu Min (Chinese Academy of Sciences, Pékin) Isabelle Rouget (UPMC, Paris) Sevket Sen (MNHN, Paris) Stanislav Štamberg (Museum of Eastern Bohemia, Hradec Králové) Paul Taylor (The Natural History Museum, Londres) COUVERTURE / COVER : Nyctereutes donnezani from Çalta, central Turkey, early Pliocene. Skull MNHN.F.ACA291, probably belonging to a male individual. Geodiversitas est indexé dans / Geodiversitas is indexed in: – Science Citation Index Expanded (SciSearch®) – ISI Alerting Services® – Current Contents® / Physical, Chemical, and Earth Sciences® – Scopus® Geodiversitas est distribué en version électronique par / Geodiversitas is distributed electronically by: – BioOne® (http://www.bioone.org) Les articles ainsi que les nouveautés nomenclaturales publiés dans Geodiversitas sont référencés par / Articles and nomenclatural novelties published in Geodiversitas are referenced by: – ZooBank® (http://zoobank.org) Geodiversitas est une revue en flux continu publiée par les Publications scientifiques du Muséum, Paris Geodiversitas is a fast track journal published by the Museum Science Press, Paris Les Publications scientifiques du Muséum publient aussi / The Museum Science Press also publish: Adansonia, Zoosystema, Anthropozoologica, European Journal of Taxonomy, Naturae, Cryptogamie sous-sections Algologie, Bryologie, Mycologie. Diffusion – Publications scientifiques Muséum national d’Histoire naturelle CP 41 – 57 rue Cuvier F-75231 Paris cedex 05 (France) Tél. : 33 (0)1 40 79 48 05 / Fax : 33 (0)1 40 79 38 40 [email protected] / http://sciencepress.mnhn.fr © Publications scientifiques du Muséum national d’Histoire naturelle, Paris, 2019 ISSN (imprimé / print) : 1280-9659/ ISSN (électronique / electronic) : 1638-9395 Phylogenetic relationships of Nyctereutes Temminck, 1838 (Canidae, Carnivora, Mammalia) from early Pliocene of Çalta, Turkey Thibault DAGUENET Sevket SEN Centre de Recherche de Paléontologie de Paris, CR2P (CNRS, MNHN, UPMC, Sorbonne Université), Département Origines et Évolution, Muséum national d’Histoire naturelle, case postale 38, 57 rue Cuvier, F-75231 Paris cedex 05 (France) [email protected] (corresponding author) Submitted on 21 December 2018 | accepted on 28 March 2019 | published on 3 October 2019 urn:lsid:zoobank.org:pub:392603E1-AC9A-4A4D-923D-307216718171 Daguenet T. & Sen S. 2019. — Phylogenetic relationships of Nyctereutes Temminck, 1838 (Canidae, Carnivora, Mammalia) from early Pliocene of Çalta, Turkey, in Bonis L. de & Werdelin L. (eds), Memorial to Stéphane Peigné: Carnivores (Hyaeno- donta and Carnivora) of the Cenozoic. Geodiversitas 41 (18): 663-677. https://doi.org/10.5252/geodiversitas2019v41a18. http://geodiversitas.com/41/18 ABSTRACT The locality of Çalta has yielded one of the richest collection of fossil canids belonging to the raccoon dog genus Nyctereutes Temminck, 1838. This locality is situated in central Anatolia and its age is determined as early Pliocene, c. 4 Ma. Ginsburg (1998) studied these canid remains and identified them as N. donnezani, which was previously known from some early Pliocene localities in southern France and Spain. Some later studies questioned this attribution. The present study provides a new description of all available material, housed in collections in Paris, Lyon and Ankara, and a detailed comparison of the cranial and dental characters with other known species of Nyctereutes. This study is completed by a cladistic analysis of cranial and dental characters to document the phylogenetic relationships of the Çalta raccoon dog. This analysis was done on the locality-based data, using the main occurrences of each species. Both morphological and cladistic analyses favour attribution of KEY WORDS Canidae, the Çalta raccoon dog to N. donnezani, in agreement with Ginsburg (1998). In addition, this study Nyctereutes, reveals that possible sexual dimorphism in the size and proportions of some cranial elements, such as Pliocene, stronger sagittal and nuchal crests, a better defined temporal line, a thicker zygomatic process of the Turkey, phylogeny, maxillary bone, a braincase that is elongated but less round and, in particular, stronger subangular sexual dimorphism. lobe in male individuals. GEODIVERSITAS • 2019 • 41 (18) © Publications scientifiques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com 663 Daguenet T. & Sen S. RÉSUMÉ Relations phylogénétiques de Nyctereutes Temminck, 1838 (Canidae, Carnivora, Mammalia) du Pliocène inférieur de Çalta, Turquie. La localité de Çalta a livré l’une des plus riches collections de canidés fossiles appartenant au genre Nyctereutes Temminck, 1838. Cette localité est située en Anatolie centrale et elle est datée du Pliocène inférieur, c. 4 Ma. Ginsburg (1998) a étudié les carnivores de Çalta et a identifié les fossiles de chien viverrin comme N. donnezani, une espèce précédemment connue dans quelques localités du Pliocène inférieur du sud de la France et de l’Espagne. Certaines études ultérieures ont remis en cause cette attribution. La présente étude fournit une nouvelle description de tout le matériel disponible, hébergé dans les collections de Paris, Lyon et Ankara, ainsi qu’une comparaison détaillée des caractères crâniens et dentaires avec d’autres espèces connues de Nyctereutes. Cette étude est complétée par une analyse cladistique de caractères crâniens et dentaires afin de documenter les relations phylogénétiques du chien viverrin de Çalta. Cette analyse a été réalisée par gisement plutôt que par espèce, en utilisant les principales occurrences de chaque espèce. Les analyses morphologiques et cladistiques sont toutes deux MOTS CLÉS Canidae, en faveur de l’attribution du chien viverrin de Çalta à N. donnezani, comme l’avait précédemment fait Nyctereutes, Ginsburg (1998). En outre, cette étude révèle qu’un dimorphisme sexuel se manifeste dans la taille et Pliocène, les proportions de certains éléments crâniens, tels que des crêtes sagittales et nuchales plus fortes, une Turquie, phylogénie, ligne temporelle mieux définie, un processus zygomatique du maxillaire plus épais, une boite crânienne dimorphisme sexuel. allongée mais moins ronde et en particulier un lobe sous-angulaire fort chez les individus mâles. INTRODUCTION to N. donnezani (Depéret, 1890), the type locality of which is Perpignan in southern France. To date, this is the unique The Old World Canidae are immigrants from North America occurrence of raccoon dog in Turkey. Ginsburg (1998) com- where the earliest fossil occurrences date back to the middle pared the Çalta fossils with N. donnezani and N. megamastoides, Eocene, whereas the oldest representatives of the family in in addition to Vulpes and Canis. However, this identification Eurasia are from the late Miocene (McKenna & Bell 1997). was not sufficiently convincing since, as we will see below, Among the representatives of this large family, the genus some later studies have questioned the assignment of the Çalta Nyctereutes Temminck, 1838 occupies a particular position in raccoon dog to N. donnezani. having a wide distribution all over the three continents of the In the neighbouring countries of Turkey, raccoon dog fossils Old World. The only extant species of this genus, the raccoon are recorded in Greece, Bulgaria and Romania. From Greece, dog N. procyonoides (Gray, 1834), is naturally restricted to East Koufos (1997) referred a skull from Megalo Emvolon to Asia, i.e., SE Russia, eastern China, Korea, Japan and north N. tingi, while the remains of N. megamastoides are reported Vietnam (Ward & Wurster-Hill 1990). It was introduced in from several localities of late Pliocene and early Pleistocene western Russia in 1920s for fur production, and individuals age (Koufos 1993; Athanassiou 1996; De Vos et al. 2002). who escaped from production centres gradually populated In Bulgaria, Spassov (1997, 2003) mentioned the presence eastern and central Europe. At present the raccoon dog is of N. cf. tingi in the early Pleistocene locality of Varshets. In known as an invasive species in Europe from western Russia Romania, Simionescu (1930, 1932) described some cranial and the Caucasus to eastern France and northern Italy, and remains as Vulpes donnezani from the early Pliocene localities from Romania to Norway (Kauhala & Kowalczyk 2011). of Malusteni and Beresti that Radulescu et al. (2003) referred Fossil species of Nyctereutes are known from Europe, Asia to Eucyon odessanus. These authors also mentioned the presence and Africa. The earliest fossil occurrence of this genus goes of
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