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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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Abstract

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.

Kurzfassung

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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Abbreviations

IVPP (V):

Institute of Vertebrate Paleontology and Paleoanthropology, Beijing, China

NMNH-A:

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

NHMW:

Naturhistorisches Museum Vienna, Austria

PMU (M):

Collection of the Evolution Museum of Uppsala University, Sweden

C:

Upper canine

c:

Lower canine

I:

Upper incisor

i:

Lower incisor

M:

Upper molar

m:

Lower molar

PM:

Upper premolar

pm:

Lower premolar

DIA:

Diastema

L:

Mesiodistal length

W:

Bucco-lingual width

H:

Crown height

PadH:

Height of the paraconid

PadL:

Length of the paraconid

PrdH:

Height of the protoconid

PrdL:

Length of the protoconid

BU:

Maximum length of the tympanic bullae

LAL:

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

LB:

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

LBO:

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

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 caudal border of the occipital

LO:

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

LP:

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

LPA:

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

LPL:

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

LPP:

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

LPPF:

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

LR:

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

LSD:

Length of the upper dental series

TFL:

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

WN:

Width of the nasals

WSU:

Skull width across buccal margins of the upper canines

WSP:

Skull width across distal buccal margins of PM4 s

CM1:

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

COM1:

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

DMS:

Depth of the mandibular symphysis distally of the lower canine

DMM:

Depth of the mandible under m1 (labially)

HRM:

Height of the mandibular ramus

LCOR:

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

LM:

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

LSY:

Length of the lower jugal series

MAM:

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

TDM:

Transverse diameter of the mandible at m1

HD:

Hadjidimovo locality, Bulgaria

LX:

Indication for localities in Linxia Basin

References

  • Alcalá, L., C. Sesé, E. Herráez, and R. Adrover. 1991. Mamíferos del Turoliense inferior de Puente Minero (Teruel, España). Boletín Real Sociedad Española de Historia Natural 86 (1–4): 205–251.

    Google Scholar 

  • Berta, A. 1987. The sabercat Smilodon gracilis from Florida and a discussion of its relationships (Mammalia, Felidae, Smilodontini). Bulletin of the Florida State Museum, Biological Science 31 (1): 1–63.

    Google Scholar 

  • Beaumont, G. 1975. Recherches sur les félidés (Mammifères, Carnivores) du Pliocène inférieur des sables à Dinotherium des environs d’Eppelsheim (Rheinhessen). Archives des Sciences, Genève 28: 369–405.

    Google Scholar 

  • Beaumont, G. 1978. Notes complémentaires sur quelques félidés (Carnivores). Archives des Sciences, Genève 31: 219–227.

    Google Scholar 

  • Bowdich, T.E. 1821. An analysis of the natural classifications of Mammalia, for the use of students and travellers, 1–115. Paris: J. Smith.

  • Clavel, J., G. Merceron, L. Hristova, N. Spassov, D. Kovachev, and G. Escarguel. 2012. On Mesopithecus habitat: insights from late Miocene fossil vertebrate localities of Bulgaria. Journal of Human Evolution 63: 162–179.

    Article  Google Scholar 

  • Deng, T. 2004. Evolution of the late Cenozoic mammalian faunas in the Linxia Basin and its background relevant to the uplift of the Qinghai-Xizang Plateau. Quaternary Sciences 24 (4): 413–420. (Chinese with English abstract).

    Google Scholar 

  • Deng, T., X.-M. Wang, X.-J. Ni, and L.-P. Liu. 2004. Cenozoic stratigraphic sequence of the Linxia Basin in Gansu, China and its evidence from mammal fossils. Vertebrata PalAsiatica 42 (1): 45–66. (Chinese and English).

    Google Scholar 

  • Deng, T. 2006. Chinese Neogene mammal biochronology. Vertebrata PalAsiatica 44 (2): 143–163.

    Google Scholar 

  • Deng, T., W. He, and S.-Q. Chen. 2008. A new species of the Late Miocene tapirs (Perissodactyla, Tapiridae) from the Linxia Basin in Gansu, China. Vertebrata PalAsiatica 46 (3): 190–209. (Chinese and English).

    Google Scholar 

  • Deng, T., Z.-X. Qiu, B.-Y. Wang, X.-M. Wang, and S.-K. Hou. 2013. Late Cenozoic biostratigraphy of the Linxia Basin, northwestern China. In Fossil mammals of Asia: neogene biostratigraphy and chronology, ed. X.-M. Wang, L.J. Flynn, and M. Fortelius, 243–273. Columbia: Columbia University Press.

    Chapter  Google Scholar 

  • Fischer von Waldheim, G. 1817. Adversaria zoologica. Mémoires de la Société Impériale des Naturalistes de Moscou 5: 357–428.

  • Geraads, D., and N. Spassov. 2009. Rhinocerotidae (Mammalia) from the late Miocene of Bulgaria. Palaeontographica 287: 99–122.

    Article  Google Scholar 

  • Gill, T. 1872. Arrangement of the families of mammals with analytical tables. Smithsonian Micellaneous Collections 11(4): 1–98.

  • Ginsburg, L. 1999. Order Carnivora. In The Miocene land mammals of Europe, eds. G.E. Rössner, and K. Heißig, 109–148. München: Pfeil-Verlag.

    Google Scholar 

  • Goloboff, P.A., J.S. Farris, and K.C. Nixon. 2008. TNT, a free program for phylogenetic analysis. Cladistics 24: 774–786.

    Article  Google Scholar 

  • GSA Rock Color Chart. 1991. The Geological Society of America Rock-Color Chart with Genuine Munsell Color Chips. USA: Munsell Color.

    Google Scholar 

  • Hou, L.H., Z.-H. Zhou, F.-C. Zhang, and Z. Wang. 2005. A Miocene ostrich fossil from Gansu Province, northwest China. Chinese Science Bulletin 50 (16): 1808–1810.

    Article  Google Scholar 

  • Hristova, L., and N. Spassov. 2005. Taxonomy and Ecology of the late Miocene Hipparions of Bulgaria: Implication to the questions of the Balkans Turolian stratigraphy and environmental changes, 24. Institute of Botany, Bulgarian Academy of Sciences, Abstracts: In Annual NECLIME Meeting.

    Google Scholar 

  • Hristova, L., D. Geraads, G. Markov, and N. Spassov. 2013. Late Miocene mammals from Kocherinovo. Southwestern Bulgaria. Acta Zoologica Bulgarica 65 (4): 517–529.

    Google Scholar 

  • Kaup, J. 1832. Vier neue Arten urweltlicher Raubthiere welche im zoologischen Museum zu Darmstadt aufbewart werden. Archiv für Mineralogie 5: 150–158.

    Google Scholar 

  • Kittl, E. 1887. Beiträge zur Kenntnis der fossilen Säugethiere von Maragha in Persien. Annalen des Kaiserlich Königlichen Naturhistorischen Hofmuseums Wien 1: 317–338.

    Google Scholar 

  • Kretzoi, N. 1929. Materialien zur phylogenetischen Klassifikation der Aeluroïdeen. X Congres International de Zoologie, Budapest 2: 1293–1355.

    Google Scholar 

  • Kretzoi, N. 1938. Die Raubtiere von Gombaszög nebst einer Übersicht der Gesamtfauna. Annales Historico-Naturales Musei Nationalis Hungarici 31: 88–157.

    Google Scholar 

  • Koufos, G., N. Spassov, and D. Kovatchev. 2003. Study of Mesopithecus (Primates, Cercopithecoidea) from the Late Miocene of Bulgaria. Palaeontographica Abteilung A 269 (1–3): 39–91.

    Google Scholar 

  • Li, Y. 2015. Restudy of Metailurus major from Yushe Basin, Shanxi Province reported by Teilhard de Chardin and Leroy. Vertebrata PalAsiatica 52 (4): 467–485.

    Google Scholar 

  • Maddison, W.P., and D.R. Maddison. 2015. Mesquite: a modular system for evolutionary analysis. Version 3.04. Available: http://mesquiteproject.org.

  • McKenna, M.C., and S.K. Bell. 1997. Classification of mammals above the species level. New York: Columbia University Press.

    Google Scholar 

  • Merceron, G., A. Zazzo, N. Spassov, D. Geraads, and D. Kovachev. 2006. Bovid paleoecology and paleoenvironments from the Late Miocene of Bulgaria: evidence from dental microwear and stable isotopes. Palaeogeography, Palaeoclimatology, Palaeoecology 241 (3–4): 637–654.

    Article  Google Scholar 

  • Merriam, J.C., and C. Stock. 1932. The Felidae of Rancho La Brea. Carnegie Institution of Washington 22: 1–231.

    Google Scholar 

  • Montoya, P. 1994. Los Macromamíferos del Mioceno superior del área de Crevillente (Alicante). Unpublished PhD thesis. Universitat de València, 421 pp.

  • Morlo, M. 1997. Die Raubtiere (Mammalia, Carnivora) aus dem Turolium von Dorn-Dürkheim 1 (Rheinhessen). Teil 1: Mustelidae, Hyaenidae, Percrocutidae. Felidae. Courier Forschungs-Institut Senckenberg 197: 11–47.

    Google Scholar 

  • Morales, J., and D. Soria. 1977. Presencia de la asociación Machairodus-Paramachairodus en Concud (Teruel). Teruel 57–58: 1–9.

    Google Scholar 

  • Morales, J. 1984. Venta del Moro: su macrofauna de mamíferos y biostratigrafía continental del Mioceno terminal mediterráneo, 1–340. Madrid: Universidad Complutense.

    Google Scholar 

  • Nikolov, I. 1973. The Genus Hipparion in Bulgaria. Ph.D. Thesis. Institute of Geology, Bulgarian Acadamy of Sciences (Bulgarian).

  • Ozansoy, F. 1965. Étude des gisements continentaux et de mammifères du Cénozoique de Turquie. Mémoire de la Societée Géologique de France (n. série) 44:1–92.

  • Nikolov, I. 1985. Catalogue of the localities of tertiary Mammals in Bulgaria. Bulgarian Academy of Sciences 21: 43–62.

    Google Scholar 

  • Pilgrim, G. 1913. The correlation of the siwaliks with Mammal Horizons of Europe. Records of the Geological Survey of India 43: 264–326.

    Google Scholar 

  • Pilgrim, G. 1931. Catalogue of the Pontian Carnivora of Europe in the Department of Geology. London: British Museum of Natural History.

  • Salesa, M.J., P. Montoya, L. Alcalá, and J. Morales. 2003. El género Paramachairodus Pilgrim, 1913 (Felidae, Machairodontinae) en el Mioceno superior Española. Coloquios de Paleontología, Volumen Extraordinario 1: 603–615.

    Google Scholar 

  • Salesa, M.J., M. Antón, A. Turner, and J. Morales. 2005a. El origen de los félidos machairodontinos: aspectos funcionales de la anatomía cráneo-cervical de Paramachairodus ogygia (Kaup, 1832) (Felidae, Machairodontinae) de Batallones-1 (Vallesiense, MN 10). Revista Española de Paleontología 20 (2): 133–141.

    Google Scholar 

  • Salesa, M.J., M. Antón, A. Turner, and J. Morales. 2005b. Aspects of the functional morphology in the cranial and cervical skeleton of the sabre-toothed cat Paramachairodus ogygia (Kaup, 1832) (Felidae, Machairodontinae) from the Late Miocene of Spain: implications for the origins of the machairodont killing bite. Zoological Journal of the Linnean Society 144: 363–377.

    Article  Google Scholar 

  • Salesa, M.J., M. Antón, A. Turner, and J. Morales. 2006. Inferred behaviour and ecology of the primitive sabre-toothed cat Paramachairodus ogygia (Felidae, Machairodontinae) from the Late Miocene of Spain. Journal of Zoology 268: 243–254.

    Article  Google Scholar 

  • Salesa, M.J., M. Antón, A. Turner, L. Alcalá, P. Montoya, and J. Morales. 2010. Systematic revision of the Late Miocene sabre-toothed felid Paramachaerodus in Spain. Palaeontology 53: 1369–1391.

    Article  Google Scholar 

  • Spassov, N. 2002. The Turolian Megafauna of West Bulgaria and the character of the Late Miocene “Pikermian biome”. Bollettino della Societa Paleontologica Italiana 41 (1): 69–81.

    Google Scholar 

  • Spassov, N., and D. Geraads. 2004. Tragoportax Pilgrim and Miotragocerus Stromer (Mammalia, Bovidae) from the Turolian of Hadjidimovo, Bulgaria, and a revision of the Late Miocene Mediterranean Boselaphini. Geodiversitas 26 (2): 339–370.

    Google Scholar 

  • Spassov, N., and D. Geraads. 2015. A new felid from the late Miocene of the Balkans and the contents of the genus Metailurus Zdsndky, 1924 (Carnivora, Felidae). Journal of Mammalian Evolution 22 (1): 45–56.

    Article  Google Scholar 

  • Tseng, Z.-J., X.-M. Wang, G.J. Slater, G.T. Takeuchi, Q. Li, J. Liu, and G.-P. Xie. 2014. Himalayan fossils of the oldest known pantherine establish ancient origin of big cats. Proceedings of the Royal Society B 281: 20132686.

    Article  Google Scholar 

  • Vatsev, M. 1980. Lithostratigraphy of the Neogene sedimentary rocks of the Gotse Delchev basin. Annual of the Higher Institute of Mine and Geology 25: 103–115.

    Google Scholar 

  • Wagner, A. 1857. Neue Beiträge zur Kenntniss der fossilen Säugthier-Ueberreste von Pikermi. Abhandlungen der Bayerischen Akademie der Wissenschaften, Mathematisch-naturwissenschaftlichen Klasse 8: 111–158.

    Google Scholar 

  • Weithoffer, K. 1888. Beiträge zur Kenntniss der Fauna von Pikermi bei Athen. Beiträge zur Paläontologie und Geologie Österreich-Ungarns und des Orients 6: 225–292.

  • Werdelin, L. 1981. The evolution of lynxes. Annales Zoologici Fennici 18: 37–71.

    Google Scholar 

  • Zdansky, O. 1924. Jungtertiäre Carnivoren Chinas. Palaeontologica Sinica 2: 1–149.

    Google Scholar 

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Acknowledgements

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). https://doi.org/10.1007/s12542-017-0371-7

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