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A New Lophocyonid, Izmirictis Cani Gen. Et Sp. Nov. (Carnivora: Mammalia)

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A New Lophocyonid, Izmirictis Cani Gen. Et Sp. Nov. (Carnivora: Mammalia) Journal of Systematic Palaeontology, 2019 Vol. 0, No. 0, 1–12, http://dx.doi.org/10.1080/14772019.2018.1529000 A new lophocyonid, Izmirictis cani gen. et sp. nov. (Carnivora: Mammalia), from the lower Miocene of Turkey a b c,dà e,f g Jorge Morales , Serdar Mayda , Alberto Valenciano , Daniel DeMiguel and Tanju Kaya aDepartamento de Paleobiologıa, Museo Nacional de Ciencias Naturales, CSIC, C/ Jose Gutierrez Abascal, 2. 28006, Madrid, Spain; bDepartment of Biology, Ege University, Bornova, 35100 Izmir, Turkey; cResearch and Exhibitions Department, Iziko Museums of South Africa, P.O. Box 61, Cape Town, South Africa; dDepartment of Biological Sciences, University of Cape Town, Palaeobiological Research Group, Private Bag X3, Rhodes Gift 7701, Cape Town, South Africa; eFundacion ARAID/Universidad de Zaragoza, Departamento de Ciencias de la Tierra, Area de Paleontologıa, Pedro Cerbuna 12, 50009 Zaragoza, Spain; fInstitut Catala de Paleontologia Miquel Crusafont, Universitat Autonoma de Barcelona. Edifici ICTA-ICP, C/ Columnes s/n, Campus de la UAB, 08193 Cerdanyola del Valles, Barcelona, Spain; gNatural History Museum, Ege University, Bornova, 35100 Izmir, Turkey (Received 31 May 2018; accepted 22 September 2018) Lophocyonids are one of the more enigmatic groups of Carnivora in the European Miocene fossil record. Lophocyonids are clearly distinguished from other Feliformia by their peculiar lophodont dental morphology. For this reason, the systematic relationships of the family have been controversial. Here we describe and interpret dental remains from the early Miocene of Sabuncubeli, Turkey, which we attribute to a new genus and species: Izmirictis cani. The phylogenetic analysis allows us to include Izmirictis within a monophyletic group, Lophocyonidae new rank, characterized by the molarization of the anterior premolars (P3 and p4), the lophodont adaptation of the molar dentition and the complex morphology of the incisors. The phylogenetic analysis shows a close relationship between Izmirictis and primitive hyaenids (represented herein by Protictitherium). The divergence between Lophocyonidae and Hyaenidae is estimated by biochronological data to have occurred during the early Miocene (MN2). Dental microwear analysis, although limited by poor dental enamel preservation, indicates that the pronounced lophodonty in Izmirictis cani could be connected to a herbivorous feeding habit. http://zoobank.org/urn:lsid:zoobank.org:pub:8D75382D-6C22-41C1-A47F-BA3EAF766881 Keywords: Lophocyonidae; Feliformia; systematics; Miocene; Europe; Turkey Introduction this species the genus Sivanasua (Aeluravus was pre- occupied by a glirid) and included in it two new species, Lophocyoninae Fejfar, Schmidt-Kittler & Zacharov, both of which were later classified as Adapidae by 1987 is one of the most peculiar groups of terrestrial Gingerich & Sahni (1984). However, the genus name is carnivorans in the fossil record due to its lophodont den- valid, and is generally considered to be close to the tition, which is particularly advanced in the most coati-like procyonid carnivorans. After the study of derived members of the group. This subfamily includes Schmidt-Kittler (1983) they were considered to be the genus Sivanasua Pilgrim, 1932 with the species S. primitive feliforms, classified in their own subfamily viverroides (Schlosser, 1916) and S. moravica Fejfar & Lophocyoninae, within the Viverridae (Fejfar Schmidt-Kittler, 1984; Euboictis aliverensis Schmidt- et al. 1987). Kittler, 1983; the genus Lophocyon Fejfar et al., 1987 The aim of the present work is the description of a with the species L. carpathicus Fejfar et al., 1987 and new form from the lower Miocene of Sabuncubeli L. paraskevaidisi Koufos et al., 1995, all of which are (Turkey) (de Bruijn et al. 2006) tentatively attributed to from the early–middle Miocene of Europe. The first this interesting group of carnivorans by Mayda (2004, interpretations of the fossils by Schlosser (1916) related 2010) and Hoek Ostende et al.(2015). The primitive them to the extant red panda Ailurus fulgens Cuvier in morphology of its dentition, in comparison with the Geoffroy Saint-Hilaire & Cuvier, 1825. Schlosser other representatives of the group, yields important clues (1916) described the first species as ‘Aeluravus’ viver- about Lophocyoninae phylogenetic relationships. It is roides on the basis of fossils from the site of Attenfeld, the first representative of the group to be found outside MN4 (Bavaria, Germany). Pilgrim (1932) proposed for of continental Europe. à Corresponding author. Email: [email protected] # The Trustees of the Natural History Museum, London 2019. All rights reserved. Published online 29 Jan 2019 2 J. Morales et al. Fossil locality Thymiana-B, Chios Island, Greece, MN5 (Koufos 2011), of which a cast is kept at the MNHN; the holotype of The locality of Sabuncubeli is situated in the lower part Kichechia zamanae Savage (1965)fromRusinga,Kenya, of the Soma Formation (Fig. 1), along the forest road lower–middle Miocene 17–15 Ma (Peppe et al. 2009) that leads from the main road between Manisa and housed at the NHMUK; Herpestides antiquus (Blainville, Bornova to the village of Sarnic, 15 km NW of Izmir 1842), from Saint-Gerand-le-Puy, France, MN2a, housed (de Bruijn et al. 2006; Kaya et al. 2007). The at the MNHN and described by Viret (1929)and Sabuncubeli micromammal association is considered to Beaumont (1967); Protictitherium crassum (Deperet, be somewhat older than the one from Kesekoy€ 1892) from the Vallesian of Spain from the sites of Los (Krijgsman et al. 1996) and it fits best into the lower Valles de Fuentiduena,~ MN9 and Cerro de los Batallones, part of MN 3. According to a preliminary identification MN10 (Fraile 2017). The taxa Lophocyon carpathicus (Mayda 2010) the carnivorans found in the site are: an Fejfar et al., 1987 from Kosice-Bankov, Slovakia, MN5, amphicyonid (Cynelos sp. nov.), a musteloid Legetetia nandii Schmidt-Kittler (1987) from Legetet, (Stromeriella sp.), a viverrid (Semigenetta elegans), a Kenya, lower Miocene 19 Ma (Werdelin 2010), and feliformia (Palaeogale sp.) and an undetermined felid Protictis spp. described in Flynn & Galiano (1982), that cannot yet be formally attributed to a genus. Gingerich & Winkler (1985) and Meehan & Wilson Besides its diverse carnivore assemblage, Sabuncubeli (2002) from the middle Paleocene of North America has also yielded a new tragulid (Dorcatherium sp. nov.), were studied based on the original publications. The an early cervid (Procervulus sp.) and a small rhino extant carnivorans studied were the viverrids Viverra tan- (Protaceratherium sp.), which are under study by SM. galunga Gray, 1832,andHemigalus derbyanus (Gray, 1837) housed at MNCN and MNHN. Measurements in mm (L ¼ length, W ¼ width) were made with Mitutoyo Absolute digital callipers accurate to 0.1 mm. P4 ¼ length Material and methods mestastyle-parastyle, width parastyle-protocone. The pho- tographs of the dentition of I. cani were made with a Institutional abbreviations Scanning Electron Microscope Fei, model Quanta 200, at EUNHM: Ege University Natural History Museum, the MNCN. Bornova, Izmir, Turkey; IPS: collection of the Institut Catala de Paleontologia Miquel Crusafont (ICP), Universitat Autonoma de Barcelona, Spain; MNCN: Cladistic analysis Museo Nacional de Ciencias Naturales, Madrid, Spain; We performed a cladistic analysis to better assess the MNHN:Museum national d’Histoire naturelle, Paris, relationships of Izmirictis with other taxa possessing France; NHMUK: Natural History Museum, London, UK. similar lophodont dentitions, such as Sivanasua viver- roides, Lophocyon spp. (L. paraskevaidisi and L. carpa- thicus) and Euboictis aliverensis, together with the Studied material primitive hyaenid Protictitherium crassum, and lower– The fossil remains of Izmirictis cani gen. et sp. nov. are middle Miocene genera with uncertain relationships housed in EUNHM: PV-54000: right I3; PV-54001: left including Herpestides antiquus, Kichechia zamanae and I2; PV-54003: right P3; PV-54004: left P4; PV-54005a: Legetetia nandii, as well as the living viverrids Viverra right M1; PV-54005b: fragment of right M1; PV-54006: tangalunga and Hemigalus derbyanus. The cladistic ana- left M2; PV-54007: right i2; PV-54008: left p1 or p2; lysis includes 11 taxa and 67 equally weighted, PV-54009: right p4; PV-54010: left m1. unordered dental and cranial characters generated by us. For comparison we used Sivanasua viverroides The character list, data matrix and nexus files are avail- (Schlosser, 1916) from Rothenstein 1/13 described by able in the Supplementary material. We selected Fejfar et al.(1997), including a cast of specimen 0030 Protictis spp. as the outgroup. The analysis was per- – ^ à stored at the MNCN, and the P4 P3 from Chene de formed using PAUP 4.0b10 (Swofford 2002). Navere, France, MN4, cast stored in the MNHN, figured by Ginsburg & Morales (1999), Sivanasua antiqua (Crusafont-Pairo, 1959) from Can Canals, Spain, lower Systematic palaeontology Miocene, MN4, housed at the ICP; the holotype of Euboictis aliverensis Schmidt-Kittler (1983)fromAliveri, Order Carnivora Bowdich, 1821 Greece, MN4 (Hoek Ostende et al. 2015), of which a Suborder Feliformia Kretzoi, 1945 cast is preserved at the MNCN; the holotype of Family Lophocyonidae Fejfar, Schmidt-Kittler & Lophocyon paraskevaidisi Koufos et al.(1995)from Zacharov, 1987 (new rank) A new lophocyonid from Turkey 3 Figure 1. A, geological map of the Izmir area, Turkey, showing the location of the Sabuncubeli mammal fossil
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