(Rodentia, Mammalia) from the Paleogene of South-East Serbia, the First Record of the Family in Europe
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Palaeobio Palaeoenv DOI 10.1007/s12549-017-0301-4 ORIGINAL PAPER On a new diatomyid (Rodentia, Mammalia) from the Paleogene of south-east Serbia, the first record of the family in Europe Zoran Marković1 & Wilma Wessels2 & Andrew A. van de Weerd2 & Hans de Bruijn2 Received: 31 March 2017 /Revised: 19 May 2017 /Accepted: 28 August 2017 # The Author(s) 2017. This article is an open access publication Abstract A new diatomyid genus and species, Inopinatia Introduction balkanica, from the early Oligocene of south-east Serbia is described, and the affinities between the Diatomyidae and During the ongoing research of the Natural History Museum Ctenodactylidae are discussed. Inopinatia balkanica nov. of Belgrade, upper Eocene and lower Oligocene non-marine gen. nov. sp. seems to have retained its deciduous teeth deposits in basins of southern Serbia were investigated for the throughout life just as all other species of the family. The only presence of fossil mammals. Early Oligocene rodent faunas other diatomyid described from outside south-east Asia which were discovered in 2010 by Zoran Marković and Miloš is Pierremus explorator López-Antoñanzas, 2010 is trans- Milivojević (Natural History Museum) in the Babušnica- ferred to the thryonomyid species Paraphiomys knolli Koritnica basin. The discovery was soon followed by López-Antoñanzas and Sen, 2005. collecting campaigns in close cooperation with Hans de Bruijn and Wilma Wessels of the University of Utrecht. At present, two rodent faunas of late Eocene age and five of early Keywords Rodentia . Diatomyidae . Oligocene . Serbia . Oligocene age have been found. For the geological setting, the Biogeography geographical position and an overview of fossil content of these sites we refer to Bruijn et al. (in press). The distribution of rodent taxa in the faunas is shown in Fig. 1. In this paper, we will describe a new genus and species of the Diatomyidae from the early Oligocene localities in the B This is the second paper in the series The Paleogene rodent faunas from Babušnica-Koritnica basin. It is the first occurrence outside south-east Serbia^. of Asia of this rodent family, and the Serbian fossils are This article is registered in Zoobank under urn:lsid:zoobank. among the oldest representatives known. Prior to the org:pub:AAF7D4B5-B475-41F7-8BE5-8123A3FB0457 findings in southern Serbia, fossils of this family were known only from the Oligocene and Miocene of Asia. The * Zoran Marković [email protected] extant diatomyid Laonastes aenigmamus Jenkins et al., 2005 was discovered in Laos in 2004. A few years later, Dawson * Wilma Wessels [email protected] et al. (2006) included this new animal in the Diatomyidae. The discovery of a new diatomyd genus in Serbia, a family * Andrew A. van de Weerd [email protected] so far know only from Asia, suggests a biogeographic pattern similar to that of the few findings of larger mammals in upper * Hans de Bruijn [email protected] Eocene and Oligocene deposits in the Balkan countries of Bulgaria and Romania. These show strong taxonomic affini- 1 Natural History Museum in Belgrade, Njegoševa 51, ties to Asian species while they are absent in the well-known Belgrade 11000, Serbia Central and Western European faunas. 2 Department of Earth Sciences, Utrecht University, Heidelberglaan 2, The Eocene-Oligocene transitional interval showed major P.O. Box 80021, 3508TA Utrecht, the Netherlands changes in climate, palaeogeography and biogeography. In Palaeobio Palaeoenv Fig. 1 Distribution of rodent species in the localities of south-east Serbia Central and Western Europe, these changes are reflected in the specimens, R—range of measurements, L—length, W— mammalian faunal turnover known as the BGrande Coupure^. width, sin—sinistral and dext—dextral. Mammal families that arrived in Central or Western Europe Abbreviations and terminology used in the description after the BGrande Coupure^ were already present in Serbia in of the microstructure of enamel are EDJ—enamel-dentine the late Eocene. However, the diatomyid family did not reach junction, HSB—Hunter-Schreger band, inclination—an- Central or Western Europe. gle between the HSB and the normal to the EDJ, PI— portio interna, PE—portio externa, OES—outer enamel surface, PLEX—external enamel layer without prisms, — — Methods IPM inter prismatic matrix, radial enamel enamel with parallel prisms that are at right angles to the EDJ and — The teeth that will be described and discussed have been col- BRLE basal ring of lamellar enamel. The fossil assem- lected by wet-screening fossiliferous matrix from Valniš, blages from southeastern Serbia are housed in the Strelac-1, Strelac-2, Strelac-3 and Raljin on a set of stable Natural History Museum in Belgrade (Serbia). A repre- sieves in the field (finest mesh used is 0.5 mm.). The residue sentative set of casts of rodents is kept in the collection obtained has been rewashed on a vibrating set of sieves of the department of Earth Sciences of Utrecht (Marković and Milivojević 2010). In order to reduce the con- University, the Netherlands. The locality codes of the centrate even further, it has been treated with diluted acetic Natural History Museum of Belgrade and abbreviations acid in the laboratory. The large amount of gypsum crystals in used for of the localities are 024 for Strelac-1 (STR-1), the concentrate from Raljin has been removed by heating the 025 for Strelac-2 (STR-2), 026 for Strelac-3 (STR-3), š residue to 150 °C for several hours. The fractions larger than 027 for Valni (VA) and 028 for Raljin (RA). 0.65 mm have been sorted under a microscope. The measure- ments of the teeth have been taken with a Leitz Ortholux measuring microscope with mechanical stage and measuring Taxonomy clocks. All specimens are figured as left ones. If the original is from the right side, this is indicated by underlining its number Introduction on the figure. Lower case letters refer to the lower dentition, upper case letters refer to the upper dentition. Abbreviations Our new genus and species of the Diatomyidae is among for measurements and descriptions are N—number of the oldest records and the first solid record outside of Palaeobio Palaeoenv Asia of the family. However, as will be discussed below, dp4 large and molariform. Deciduous teeth are retained Inopinatia balkanica nov.gen.nov.sp.sharessomeba- throughout life. DP4, M1 and M2 with three roots. sic dental characters with the Ctenodactylidae. Enterostyle absent in the upper cheek teeth. The m1 and m2 The reconstruction of the dentition of rodents with pre- have three roots, two roots anteriorly and one posteriorly, a molars on the basis of isolated teeth often introduces un- very large hypoconulid and a small ectostylid. The dp4 has an certainties in recognising the tooth position in the jaw ectostylid behind the hypoconid. Microstructure of the enamel (Flynn et al. 1986). In the case of Inopinatia nov. gen., of the incisors multiserial. Molar enamel is S-type. Upper this problem is pertinent, because the genus closest in incisor without sulcus. dental morphology is the diatomyid Fallomus, a genus Differential diagnosis: Inopinatia differs from Fallomus and containing three species that are all exclusively known Marymus in having M3/m3 that are larger instead of smaller by isolated teeth. The terminology of the cusps in the than the M2/m2 and in retaining the endoloph of the DP4 and descriptions below is shown in Fig. 2. the ectolophid of the m1, m2 and m3. In Inopinatia, the large hypoconulid is the highest cusp in unworn dp4, m1, m2 and is Inopinatia nov. gen. never reduced as in some m1 and m2 of Fallomus and Marymus. Type species: Inopinatia balkanica nov. sp. Type locality: Valniš Inopinatia balkanica nov. sp. Age: Early Oligocene (Fig. 3a–h, Fig. 4a–h) Derivation nominis: from the Latin word inopinatus meaning unexpected Holotype: m1 dext. (Fig. 4d, no. VA-213) Diagnosis: Inopinatia is a large diatomyid with an incipiently Type locality: Valniš bi-lophodont pattern of the cheek teeth. Endoloph in upper Age: Early Oligocene molars absent, ectolophid in lower molars weak. DP4 and Material and measurements: see Tables 1 and 2. Fig. 2 Cusp terminology for the cheek teeth of Inopinatia nov. gen. Palaeobio Palaeoenv Fig. 3 Upper cheek teeth of Inopinatia balkanica nov. sp. from Valniš. a DP4 (VA-108). b DP4 (VA-115). c M1 (VA-143). d M1 (VA-152). e M2 (VA- 142). f M2 (VA-152). g M3 (VA-161). h M3 VA-163. Fallomus razae from its type locality Pazbogi Nala GSP-417. i DP4. j M1. k M2. l M3 Description of the upper cheek teeth (Fig. 3a–h) metaloph that is directed towards the protocone, is clearly distinguishable from the posteroloph in fresh specimens. The The DP4 is a molariform tooth. The short anteroloph connects posteroloph connects the hypocone with the metacone in most the labial side of the protocone with the anterior side of the specimens, but there may be a narrow notch that isolates the paracone in most DP4, but in some others, it is very short and metacone. The M1 and M2 have three roots each. developed as a parastyle. There is no wear facet on the anterior The M3 is large, and the morphology of the M3 is similar to side, so Inopinatia did not have a DP3. The protocone and the that of the M1 and M2, but compared to the M1-M2, the paracone are situated very close to one another and are fused at anteroloph is somewhat stronger; the metacone and hypocone an early wear stage. The hypocone and the metacone, the are situated closer together, and the incomplete metaloph is highest cusps in fresh specimens, are connected by the more clearly directed towards the protocone.