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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

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New Machairodontinae material from the late Miocene localities of Hezheng (China) and Hadjidimovo (Bulgaria) represents a new species of Paramachaerodus Pilgrim. Both localities are similar in age and suggest that the new species had a very large geographic range extending from northwestern China adjacent to the Tibetan Plateau (Gansu Province) to southeastern Europe or probably to all of southern Europe. The new species—Paramachaerodus transasiaticus sp. nov is characterized by a combination of features of “Promegantereon” and Paramachaerodus. This specific morphology, as well as the age of the Hezheng and Hadjidimovo (early Turolian, after the European Land Mammal Ages) put the new species in intermediary position between “Promegantereon” and Paramachaerodus. The new felid material give grounds to discuss and revise in a new light the systematic and evolution of the “Promegantereon”–Paramachaerodus lineage, which should represent successive stages of one and the same genus: Paramachaerodus Pilgrim.


Neues Material von Machairodontinae aus den obermiozänen Fundstellen Hezheng (China) und Hadjidimovo (Bulgarien) repräsentiert eine neue Art, die der Gattung Paramachaerodus Pilgrim zugeordnet werden kann. Die beiden Fundstellen sind altersgleich und deuten darauf hin, dass die neue Art ein sehr ausgedehntes Areal von Nordwest-China, im benachbarten Hochland von Tibet (Provinz Gansu), bis Südost-Europa oder möglicherweise auch ganz Südeuropa besiedelt hat. Die neue Art – Paramachaerodus transasiaticus sp. nov. – wird durch eine Kombination der Merkmale von Promegantereon und Paramachaerodus charakterisiert. Sowohl die spezifische Morphologie als auch das Alter von Hezheng und Hadjidimovo (Unteres Turolium, nach der europäischen Landsäugetier-Chronologie) stellen die neue Art in eine Zwischenposition zwischen Promegantereon und Paramachaerodus. Das neue Feliden-Material bietet eine Diskussionsgrundlage und rückt die Systematik und Evolution der Abstammungsgruppe PromegantereonParamachaerodus in ein anderes Licht, die nachfolgende Stadien von ein und derselben Gattung (Paramachaerodus Pilgrim) repräsentieren soll.

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Institute of Vertebrate Paleontology and Paleoanthropology, Beijing, China


Paleontological museum-Assenovgrad, branch of the National Museum of Natural History—Sofia, Bulgaria


Naturhistorisches Museum Vienna, Austria

PMU (M):

Collection of the Evolution Museum of Uppsala University, Sweden


Upper canine


Lower canine


Upper incisor


Lower incisor


Upper molar


Lower molar


Upper premolar


Lower premolar




Mesiodistal length


Bucco-lingual width


Crown height


Height of the paraconid


Length of the paraconid


Height of the protoconid


Length of the protoconid


Maximum length of the tympanic bullae


Length from the rostral border of the lacrimal bone to the rostral border of the premaxilla


Length from the caudal border of the foramen magnum to the rostral border of the premaxilla


Dorsal length from the caudal border of the occipital to the rostral border of the sagittal crest


Maximal dorsal length


Length from the rostral border of the sagittal crest to the caudal border of the nasals


Length from the caudal border of the nasals to the caudal border of the occipital


Length from the rostral border of the lacrimal bone to the postorbital process of the zygomatic


Length from the rostral border of the lacrimal bone to the postorbital process of the frontal bone


Length from the rostral border of the premaxilla to the caudal border of the palate


Length from the caudal border of the occipital to the rostral border of the lacrimal bone


Length from the caudal border of the palate to the caudal border of the foramen magnum


Length from the caudal border of the occipital to the rostral border of the postorbital process of the frontal bone


Length from the caudal border of the nasals to the rostral border of the premaxilla


Length of the upper dental series


Length from the postorbital process of the frontal bone to the caudal border of the occipital


Width of the nasals


Skull width across buccal margins of the upper canines


Skull width across distal buccal margins of PM4 s


Length from the mesial border of the lower canine to the distal border of m1


Length from the carnassial notch of m1 to the caudal border of the mandibular condyle


Depth of the mandibular symphysis distally of the lower canine


Depth of the mandible under m1 (labially)


Height of the mandibular ramus


Length from the mesial border of the incisors to the caudal border of the coronoid process


Length from the rostral border of the mandibular symphysis to the caudal border of the mandibular condyle


Length of the lower jugal series


Height from the dorsal border of the mandibular condyle to the central border of the angular process


Transverse diameter of the mandible at m1


Hadjidimovo locality, Bulgaria


Indication for localities in Linxia Basin


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We (especially N.S.) are very grateful to Manuel Salesa for the photos and data, and to Jorge Morales for access to the collections in the National Museum of Natural Sciences, Madrid. We (especially Y. L.) are very grateful also to Zhanxiang Qiu (IVPP) for providing the Chinese specimen and for his advises in this research, to Tao Deng (IVPP) for funding support for the cooperation work, to Shiqi Wang and Qinqin Shi (IVPP) for their help in the field work, to Thomas Stidham (IVPP) for his help in English improving and his useful comments, to Guo Xiaocong (IVPP) for her help in drawing of some pictures and Gao Wei (IVPP) for photographing. We are indebted to Velizar Simeonovski (Field Museum, Chicago and National Museum of Natural History, Sofia) who agreed to prepare the reconstruction of the life appearance of the new species described. The detailed comments by the reviewers Michael Morlo, an anonymous reviewer, and the editor Mike Reich much improved our manuscript. This work was supported by the National Natural Science Foundation of China (Grant Number 41430102), the Strategic Priority Research Program of the Chinese Academy of Sciences (XDB03020104), and the Ministry of Science and Technology of China (2012CB821906).

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Correspondence to Nikolai Spassov.

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Handling editor: Irina Ruf.

Appendix: Character analysis

Appendix: Character analysis

Definition of characters used in the cladogram (after Salesa et al. 2010) with some additions and modifications:

  1. 1.

    Nasal bones shape. 0, close to rectangular, relatively narrow. 1, sub-rectangular. 2, rectangular, relatively wide.

  2. 2.

    Shape of upper canine crown. 0, moderately flattened. 1, markedly flattened.

  3. 3.

    Upper canine crenulations. 0, absent. 1, thin crenulations. 2, well-marked crenulations.

  4. 4.

    Orientation of PM3 and PM4. 0, aligned. 1, not aligned.

  5. 5.

    Mesial cusp of PM3. 0, absent. 1, present.

  6. 6.

    PM3, disto-lingual basal expansion of crown. 0, present. 1, absent.

  7. 7.

    PM3, mesio-distal elongation of crown. 0, absent. 1, present.

  8. 8.

    Relative size PM4-PM3. 0, PM4/PM3 index ≤1.50. 1, PM4/PM3 index >1.50.

  9. 9.

    Place of PM4 protocone. 0, protocone of PM4 situated close to the mesial edge of the parastyle. 1, situated backwards.

  10. 10.

    Buccal border of PM4. 0, sinusoidal. 1, sinusoidal with strongly expanded mesial-labial edge. 2, straight.

  11. 11.

    Protocone of PM4. 0, big and elongated mesiodistally. 1, moderately elongated mesiodistally. 2, reduced.

  12. 12.

    PM4 ectostyle. 0, absent. 1, rudimentary one. 2, well-marked.

  13. 13.

    M1 crown. 0, bucco-lingually elongated. 1, shortened (rounded).

  14. 14.

    Mandibular symphysis outline. 0, without flange and without mental crest. 1, without flange and with smooth mental crest. 2, verticalized with incipient mandible flange and marked crest.

  15. 15.

    pm2. 0, present. 1, present but very rudimentary. 2, absent.

  16. 16.

    pm4 mesio-buccal basal expansion in the crown. 0, absent. 1, present.

  17. 17.

    m1 talonid. 0, elongated with very big metaconid.1, elongated with morderate metaconid. 2, very elongated with morderate metaconid (we regard this as an apomorphy as secondary cutting edge). 3, short with morderate metaconid. 4, short without metaconid.

  18. 18.

    Sagittal crest. 0, strongly prominent. 1, moderately developed.

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Li, Y., Spassov, N. 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. PalZ 91, 409–426 (2017).

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