Original article Percrocuta miocenica (Percrocutidae, Carnivora) from the middle Miocene of Brajkovac (Central Serbia) q Predrag Radovic ´ a,b,⇑,1 , Serdar Mayda c,1 , Sanja Alaburic ´ d , Zoran Markovic ´ d a Department of Archaeology, Faculty of Philosophy, University of Belgrade, 18-20 C ˇ ika Ljubina, 11000 Belgrade, Serbia b National Museum Kraljevo, 2 Trg Svetog Save, 36000 Kraljevo, Serbia c Ege Universitesi, Faculty of Science, Biology Department, 35100 Bornova-Izmir, Turkey d Natural History Museum, 51 Njegoševa, 11000 Belgrade, Serbia article info Article history: Received 15 November 2019 Accepted 8 February 2021 Available online 3 April 2021 Keywords: Percrocuta Carnivora Middle Miocene MN6 Brajkovac Serbia abstract This paper describes a well-preserved hemimandible of Percrocuta miocenica from Brajkovac (Lazarevac municipality, Central Serbia). Based on the faunal remains recovered so far, the site is attributed to Mammal Zone MN6 (Langhian, middle Miocene). Apart from Prebreza, this is only the second site where remains of P. miocenica have been found in Serbia, and the third one in the Balkans. The newly discovered fossil, along with other taxa (e.g., Bunolistriodon meidamon, Giraffokeryx punjabiensis, Alloptox sp.), under- lines the similarities between the middle Miocene Balkan and Anatolian mammal faunas. Ó 2021 Elsevier Masson SAS. All rights reserved. 1. Introduction The early to middle Miocene large mammal faunas of the Bal- kans are rare and poorly documented. The locality of Prebreza in southern Serbia is the only one providing a rich middle Miocene fauna and it has received considerable attention in the past, partic- ularly due to the fact that it represents the type locality of the peculiar medium-sized percrocutid ‘‘Percrocuta miocenica” (Pavlovic ´ and Thenius, 1965; Stefanovic ´, 2003). The Percrocutidae are represented in the Miocene extinct fau- nas of the Old World by two genera: Percrocuta and Dinocrocuta. Of these, the genus Percrocuta, a durophagous bone-cracker, is recorded from at least 20 middle Miocene localities in the Old World (NOW Community, 2019). Interpreting the systematic rela- tionships among the members of percrocutids and hyaenids has always been problematic. Until Kurtén’s (1957) detailed study on fossil hyaenids, there have not been any comments and criticism of the work of Kretzoi (1938), who formally erected the genus Per- crocuta mainly to include the small percrocutid forms from India. Kurtén (1957) resurrected Percrocuta as a subgenus of Crocuta (based on the morphology of P4) and synonymized Adcrocuta (a true hyaenid) with Percrocuta, but he did not retain the enigmatic European genus Allohyaena. Soon thereafter, Thenius (1966) erected the subgenus Percrocuta to a full generic rank in his brief study on hyaenid phylogeny. Ficcarelli and Torre (1970) subdi- vided the fossil hyaenid forms into two genera, Percrocuta Kretzoi, 1938, and Adcrocuta Kretzoi, 1938, being distinguished by the degree of reduction of the protocone in P4, although they added ‘‘percrocutoid” forms (P. tungurensis) to the genus Adcrocuta. Surprisingly, Percrocuta could also be regarded as a subgenus of Pachycrocuta according to the authors. In his detailed study on Turkish Neogene carnivores, Schmidt-Kittler (1976) confirmed the distinctiveness of Percrocuta Kretzoi, 1938 and Adcrocuta Kretzoi, 1938, mainly on the basis of the premolar and M1 mor- phologies. The author also divided the genus Percrocuta into the subgenera Percrocuta and Dinocrocuta (a new subgenus), consider- ing the ratio of P4/P3 length as a differential character. Soria (1980) also supported the taxonomic validity of Percrocuta and Adcrocuta while describing the first percrocutid from Western Europe: ‘‘Dinocrocuta gigantea” from the Spanish localities of Algezares and Ademuz. It should be noted, however, that the specimens from both localities are significantly smaller than D. gigantea and resem- ble Dinocrocuta algeriensis in size and premolar morphology; the other percrocutid, attributed to Dinocrocuta minor by Soria (1980), is also problematic. Howell and Petter (1985) provided https://doi.org/10.1016/j.geobios.2021.02.001 0016-6995/Ó 2021 Elsevier Masson SAS. All rights reserved. q Corresponding editor: Gildas Merceron. ⇑ Corresponding author at: Department of Archaeology, Faculty of Philosophy, University of Belgrade, 18-20 C ˇ ika Ljubina, 11000 Belgrade, Serbia. E-mail address: pedja_radovic@yahoo.com (P. Radovic ´). 1 These authors contributed equally to this work. Geobios 65 (2021) 41–49 Contents lists available at ScienceDirect Geobios journal homepage: www.elsevier.com/locate/geobio