Article: Systematic revision of the Late Miocene sabre-toothed felid Paramachaerodus in Spain

A systematic revision of the sabre-toothed cat genus Paramachaerodus Pilgrim, 1913 is presented. Two species are recognized within Paramachaerodus, Pa. orientalis, and Pa. maximiliani, and the genus Promegantereon Kretzoi, 1938 is retrieved to include Promegantereon ogygia. Material from the Turolian Spanish localities of Crevillente-2 (MN 11, Alicante) and Las Casiones (MN 13, Teruel), which was previously assigned to Paramachaerodus, is now included in the tribe Metailurini. The exceptional discoveries at the Spanish Vallesian (MN 10, Madrid) fossil site of Batallones-1 have made it possible to characterize the dentition and cranial anatomy of a previously very poorly known machairodontine cat, formerly included in Paramachaerodus as Pa. ogygia, which now can be distinguished from Pa. orientalis and Pa. maximiliani by the following features: canines without crenulations, P3 with a marked disto-lingual expansion, P4 without ectostyle and with a well-developed protocone, M1 bucco-lingually elongated and double-rooted, m1 with a larger talonid, and primitive mandible morphology. Thus, the population from Batallones-1 constitutes a clearly different form from the genus Paramachaerodus, and we propose its inclusion in the genus Promegantereon Kretzoi, 1938, together with an upper canine from Crevillente-2 (MN 11), very similar to those from Batallones-1. In contrast, Pa. orientalis shows the following apomorphies: crenulated canines, P3 reduced in size and without disto-lingual expansion, P4 with a clear ectostyle as well as a reduced, backwardly displaced protocone and with a rounded and single-rooted M1. The species Pa. maximiliani is characterized by its slightly larger size, crenulated canines, very elongated P3 with a moderate disto-lingual expansion and P4 and M1 similar to those of Pa. orientalis. Paramachaerodus orientalis is recorded at Puente Minero (MN 11, Teruel), Concud (MN 12, Teruel), Crevillente-15, and Crevillente-16 (both MN 12, Alicante), and Paramachaerodus maximiliani in Venta del Moro (MN 13, Valencia). The available data suggest that Pr. ogygia was present in the Iberian Late Vallesian and Early Turolian faunas (MN 10 and MN 11) but disappeared after that age. Paramachaerodus was present in the faunas throughout the Turolian, with the species Pa. orientalis and Pa. maximiliani, this latter being probably part of the same immigration event that occurred in the Late Turolian and involved other mammal taxa such as camelids and ursids.