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A New Species of Paramachaerodus (Mammalia, Carnivora, Felidae

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A New Species of Paramachaerodus (Mammalia, Carnivora, Felidae PalZ (2017) 91:409–426 DOI 10.1007/s12542-017-0371-7 RESEARCH PAPER A new species of Paramachaerodus (Mammalia, Carnivora, Felidae) from the late Miocene of China and Bulgaria, and revision of Promegantereon Kretzoi, 1938 and Paramachaerodus Pilgrim, 1913 1,2 3 Yu Li • Nikolai Spassov Received: 17 March 2016 / Accepted: 10 June 2017 / Published online: 18 August 2017 Ó Pala¨ontologische Gesellschaft 2017 Abstract New Machairodontinae material from the late Keywords Machairodontinae Á Paramachaerodus Miocene localities of Hezheng (China) and Hadjidimovo transasiaticus sp. nov. Á Promegantereon Á Late Miocene Á (Bulgaria) represents a new species of Paramachaerodus China Á Bulgaria Pilgrim. Both localities are similar in age and suggest that the new species had a very large geographic range Kurzfassung Neues Material von Machairodontinae aus extending from northwestern China adjacent to the Tibetan den obermioza¨nen Fundstellen Hezheng (China) und Plateau (Gansu Province) to southeastern Europe or prob- Hadjidimovo (Bulgarien) repra¨sentiert eine neue Art, die ably to all of southern Europe. The new species—Para- der Gattung Paramachaerodus Pilgrim zugeordnet werden machaerodus transasiaticus sp. nov is characterized by a kann. Die beiden Fundstellen sind altersgleich und deuten combination of features of ‘‘Promegantereon’’ and Para- darauf hin, dass die neue Art ein sehr ausgedehntes Areal machaerodus. This specific morphology, as well as the age von Nordwest-China, im benachbarten Hochland von Tibet of the Hezheng and Hadjidimovo (early Turolian, after the (Provinz Gansu), bis Su¨dost-Europa oder mo¨glicherweise European Land Mammal Ages) put the new species in auch ganz Su¨deuropa besiedelt hat. Die neue Art – Para- intermediary position between ‘‘Promegantereon’’ and machaerodus transasiaticus sp. nov. – wird durch eine Paramachaerodus. The new felid material give grounds to Kombination der Merkmale von Promegantereon und discuss and revise in a new light the systematic and evo- Paramachaerodus charakterisiert. Sowohl die spezifische lution of the ‘‘Promegantereon’’–Paramachaerodus lin- Morphologie als auch das Alter von Hezheng und Hadji- eage, which should represent successive stages of one and dimovo (Unteres Turolium, nach der europa¨ischen the same genus: Paramachaerodus Pilgrim. Landsa¨ugetier-Chronologie) stellen die neue Art in eine Zwischenposition zwischen Promegantereon und Para- machaerodus. Das neue Feliden-Material bietet eine Dis- kussionsgrundlage und ru¨ckt die Systematik und Evolution Handling editor: Irina Ruf. der Abstammungsgruppe Promegantereon–Paramachae- rodus in ein anderes Licht, die nachfolgende Stadien von & Nikolai Spassov ein und derselben Gattung (Paramachaerodus Pilgrim) [email protected] repra¨sentieren soll. Yu Li [email protected] Schlu¨sselwo¨rter Machairodontinae Á Paramachaerodus transasiaticus sp. nov. Promegantereon spa¨tes Mioza¨n 1 Key Laboratory of Vertebrate Evolution and Human Origins, Á Á Á Institute of Vertebrate Paleontology and Paleoanthropology, China Á Bulgarien Chinese Academy of Sciences, Beijing 100044, China 2 University of Chinese Academy of Sciences, Beijing 100049, Institutional abbreviations China IVPP (V) Institute of Vertebrate Paleontology and 3 National Museum of Natural History, Bulgarian Academy of Paleoanthropology, Beijing, China Sciences, 1, Blvd. Tzar Osvoboditel, 1000 Sofia, Bulgaria 123 410 Y. Li, N. Spassov NMNH-A Paleontological museum-Assenovgrad, LR Length from the caudal border of the nasals to the branch of the National Museum of Natural rostral border of the premaxilla History—Sofia, Bulgaria LSD Length of the upper dental series NHMW Naturhistorisches Museum Vienna, Austria TFL Length from the postorbital process of the frontal PMU (M) Collection of the Evolution Museum of bone to the caudal border of the occipital Uppsala University, Sweden WN Width of the nasals WSU Skull width across buccal margins of the upper Anatomical abbreviations canines C Upper canine WSP Skull width across distal buccal margins of PM4 s c Lower canine I Upper incisor Measurement abbreviations i Lower incisor Mandible M Upper molar CM1 Length from the mesial border of the lower m Lower molar canine to the distal border of m1 PM Upper premolar COM1 Length from the carnassial notch of m1 to the pm Lower premolar caudal border of the mandibular condyle DIA Diastema DMS Depth of the mandibular symphysis distally of L Mesiodistal length the lower canine W Bucco-lingual width DMM Depth of the mandible under m1 (labially) H Crown height HRM Height of the mandibular ramus PadH Height of the paraconid LCOR Length from the mesial border of the incisors to PadL Length of the paraconid the caudal border of the coronoid process PrdH Height of the protoconid LM Length from the rostral border of the mandibular PrdL Length of the protoconid symphysis to the caudal border of the mandibular condyle Measurement abbreviations LSY Length of the lower jugal series Cranium MAM Height from the dorsal border of the mandibular BU Maximum length of the tympanic bullae condyle to the central border of the angular LAL Length from the rostral border of the lacrimal bone process to the rostral border of the premaxilla TDM Transverse diameter of the mandible at m1 LB Length from the caudal border of the foramen magnum to the rostral border of the premaxilla Other abbreviations LBO Dorsal length from the caudal border of the HD Hadjidimovo locality, Bulgaria occipital to the rostral border of the sagittal crest LX Indication for localities in Linxia Basin LDT Maximal dorsal length LNB Length from the rostral border of the sagittal crest to the caudal border of the nasals LNO Length from the caudal border of the nasals to the Introduction caudal border of the occipital LO Length from the rostral border of the lacrimal bone Paramachaerodus Pilgrim (1913) is a machairodontine to the postorbital process of the zygomatic felid. That genus and its close relatives were poorly known LP Length from the rostral border of the lacrimal bone until the discovery of the Batallones-1 locality (Spain) with to the postorbital process of the frontal bone its abundant material (Salesa et al. 2003). Until the revision LPA Length from the rostral border of the premaxilla to of the genus, stimulated by the discovery of the important the caudal border of the palate material from Batallones-1 (Salesa et al. 2010), two, and LPL Length from the caudal border of the occipital to eventually three, Paramachaerodus species were accepted the rostral border of the lacrimal bone in the more recent taxonomic studies as valid for this LPP Length from the caudal border of the palate to the genus: Pa. ogygia from the Vallesian, Pa. orientalis from caudal border of the foramen magnum the early to middle Turolian and (eventually) Pa. maxim- LPPF Length from the caudal border of the occipital to iliani from the late Turolian (Beaumont 1975; Morales the rostral border of the postorbital process of the 1984; Morlo 1997; Ginsburg 1999; Salesa et al. 2003, and frontal bone references therein). The study of the rich fossil material 123 A new species of Paramachaerodus (Mammalia, Carnivora, Felidae) 411 from Batallones-1 led to the concept of the existence of two excavated by D. Kovachev in the 1980s, came from the genera with close features—Promegantereon Kretzoi Bulgarian Turolian site of Hadjidimovo-1 (Fig. 1b), and is known mostly in Vallesian time and the more evolved stored in the NMNH-A. Paramachaerodus Pilgrim, which replace Promegantereon The finds were compared to other similar fossils and in in Turolian (Salesa et al. 2010). Thus, Promegantereon particular to the species of Paramachaerodus and Kretzoi, considered at the time of this study as a synonym Promegantereon, discussed in the comprehensive work of of Paramachaerodus was resurrected. New material from Salesa et al. (2010). Comparisons with fossil specimens are Hezheng district (Gansu Province, China) and Hadjidi- based on direct examination of the specimens, on fig- movo (South Bulgaria) give the opportunity to develop ures and descriptions from the literature, and on excellent further ideas about the taxonomy and evolution of the photos kindly provided by M. Salesa. primitive Promegantereon–Paramachaerodus saber- The morphological descriptions follow the terminology toothed felid lineage. used by Merriam and Stock (1932), Salesa et al. (2010), and Berta (1987) with some modifications (Fig. 2a). Measurements (in mm) (Fig. 2b; Tables 1, 2, 3) were taken Materials and methods with a digital caliper. For the cladistic analysis, 18 cranio-mandibular char- The Chinese specimens (a skull and a semimandible from acters (Appendix) were scored across six taxa (Proailurus two different individuals) are collected from the Chinese lemanensis,‘‘Promegantereon’’ ogygia, Paramachaerodus late Miocene site of Shilei locality (Fig. 1a) and housed in orientalis, Paramachaerodus maximiliani, Paramachaero- the IVPP. The Bulgarian material (two partial mandibular dus transasiaticus sp. nov (Hadjidimovo material), and Pa. rami, a separate pm3, and a separate upper canine) was transasiaticus sp. nov (Hezheng material), using Proailu- rus lemanensis as an outgroup. Most of the characters analyzed are modified from the revision of the genus Paramachaerodus by Salesa et al. (2010). Supplemental characters reflect some additional features, which we consider in our taxonomical approach as important in the evolution of the mentioned taxa. The data matrix was compiled in Mesquite version 3.04 (Maddison and Mad- dison
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