Ailuridae

Ailuridae Temporal range: Oligocene–Recent PreꞒ Ꞓ O S D C P T J K Pg N
Red panda (Ailurus fulgens)
Skull and life restoration of Simocyon
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Carnivora
Superfamily:	Musteloidea
Family:	Ailuridae Gray, 1843
Genera
†Amphictinae? Amphictis Bonisictis †Magerictinae Magerictis Rothictis †Simocyoninae Actiocyon Alopecocyon Simocyon Ailurinae † Parailurus † Pristinailurus Ailurus
Extant red panda distribution

Ailuridae is a family in the mammal order Carnivora. The family consists of the red panda (the sole living representative) and its extinct relatives.

Georges Cuvier first described Ailurus as belonging to the raccoon family in 1825; this classification has been controversial ever since. ^[1] It was classified in the raccoon family because of morphological similarities of the head, colored ringed tail, and other morphological and ecological characteristics. Somewhat later, it was assigned to the bear family. ^[2]

Molecular phylogenetic studies had shown that, as an ancient species in the order Carnivora, the red panda is relatively close to the American raccoon and may be either a monotypic family or a subfamily within the procyonid family. ^{[1] [3] [4]} An in-depth mitochondrial DNA population analysis study stated: "According to the fossil record, the Red Panda diverged from its common ancestor with bears about 40 million years ago." ^{[1] [5]} With this divergence, by comparing the sequence difference between the red panda and the raccoon, the observed mutation rate for the red panda was calculated to be on the order of 10⁹, which is apparently an underestimate compared with the average rate in mammals. ^[6] This underestimation is probably due to multiple recurrent mutations as the divergence between the red panda and the raccoon is extremely deep.^{[ citation needed ]}

The most recent molecular-systematic DNA research places the red panda into its own independent family, Ailuridae. Ailuridae are, in turn, part of a trichotomy within the broad superfamily Musteloidea ^[7] that also includes the Procyonidae (raccoons), the Mephitidae (skunks), and Mustelidae (weasels); but it is not a bear (Ursidae). ^[8]

Ailurids appear to have originated during the Late Oligocene to Early Miocene in Europe. The earliest known member, Amphictis , was likely an unspecialised carnivore, based on its dentition. Ailurids subsequently dispersed into Asia and North America. The puma-sized Simocyon found in Middle Miocene-Early Pliocene of Europe, North America and China was likely a hypercarnivore. Like modern red panda it had a "false thumb" to aid in climbing. Members of the subfamily Ailurinae, which includes the modern red panda as well as the extinct genera Pristinailurus and Parailurus , developed a specialised dental morphology with blunted cusps, creating an effective grinding surface to process plant material. ^[9]

Classification

The relationship of the Ailuridae with other carnivorans is shown in the following phylogenetic tree, which is based on the molecular phylogenetic analysis of six genes in Flynn (2005), ^[10] with the musteloids updated following the multigene analysis of Law et al. (2018). ^[11]

Carnivora	Feliformia Caniformia Canidae Arctoidea Ursidae Pinnipedia Musteloidea Mephitidae Ailuridae Procyonidae Mustelidae

In addition to Ailurus, the family Ailuridae includes seven extinct genera, most of which are assigned to three subfamilies: Amphictinae, Simocyoninae, and Ailurinae. ^{[12] [13] [14] [15] [16]}

• Family Ailuridae J.E. Gray, 1843
  • Subfamily †Amphictinae  ?Winge, 1896
    • † Amphictis  ?Pomel, 1853
      • †Amphictis borbonicaViret, 1929
      • †Amphictis ambigua(Gervais, 1872)
      • †Amphictis milloquensis(Helbing, 1936)
      • †Amphictis antiqua(de Blainville, 1842)
      • †Amphictis schlosseriHeizmann & Morlo, 1994
      • †Amphictis prolongataMorlo, 1996
      • †Amphictis wintershofensisRoth, 1994
      • †Amphictis cuspidaNagel, 2003
      • †Amphictis timucuaJ.A. Baskin, 2017 ^[17]
  • Subfamily †Simocyoninae Dawkins, 1868
    • † Actiocyon Stock, 1947
      • †Actiocyon parverratisSmith et al., 2016 ^[18]
      • †Actiocyon leardiStock, 1947
    • † Alopecocyon Camp & Vanderhoof, 1940
      • †Alopecocyon gettiMein, 1958
      • †Alopecocyon goeriachensis(Toula, 1884)
    • † Protursus Crusafont & Kurtén, 1976
      • †Protursus simpsoniCrusafont & Kurtén, 1976
    • † Simocyon Wagner, 1858
      • †Simocyon primigenius(Roth & Wagner, 1854)
      • †Simocyon diaphorus(Kaup, 1832)
      • †Simocyon batalleriViret, 1929
      • †Simocyon hungaricusKadic & Kretzoi, 1927
  • Subfamily Ailurinae J.E. Gray, 1843
    • † Magerictis Ginsburg et al., 1997
      • †Magerictis imperialensisGinsburg et al., 1997
    • Tribe Pristinailurini Wallace & Lyon, 2022
      • † Pristinailurus Wallace & Wang, 2004
        • †Pristinailurus bristoliWallace & Wang, 2004
      • † Parailurus Schlosser, 1899
        • †Parailurus anglicus(Dawkins, 1888) [Parailurus hungaricusKormos, 1935]
        • †Parailurus tedfordiWallace & Lyon, 2022
        • †Parailurus baikalicusSotnikova, 2008
    • Tribe Ailurini
      • Ailurus F. Cuvier, 1825
        • Ailurus fulgens - Red panda
          • Ailurus fulgens styaniThomas, 1902 – Eastern red panda
          • Ailurus fulgens fulgensF. Cuvier, 1825 – Western red panda

In 2025, a comprehensive study describing new material of Magerictis imperialensis also reviewed the phylogeny of the entire family; it placed Magerictis and the new genus Rothictis in the new subfamily Magerictinae, recovered Simocyoninae as paraphyletic to the Ailurinae, and resurrected the family Amphictidae for Amphictis and the new genus Bonisictis . ^[19]

• Family †Amphictidae Winge, 1895
  • AmphictisPomel, 1853
    • Amphictis antiquusPomel, 1853
    • Amphictis schlosseriHeizmann & Morlo, 1994
    • The remaining species A. borbonica, A. ageniensis, A. cuspida, A. timicua were stated to be of indeterminate generic status.
  • Bonisictis
    • Bonisictis ambiguusGervais, 1872
    • Bonisictis milloquensis(Helbing, 1928)
• Family Ailuridae Gray, 1843
  • Subfamily †Magerictinae
    • MagerictisGinsburg, Morales, Soria, and Herráez, 1997
      • Magerictis imperialensisGinsburg, Morales, Soria, and Herráez, 1997
    • Rothictis
      • Rothictis prolongata(Morlo, 1996)
      • Rothictis wintershofensis(Heizmann and Morlo, 1994)
  • Subfamily †Simocyoninae Dawkins, 1868
    • ActiocyonStock, 1947
      • Actiocyon leardiStock, 1947
      • Actiocyon parverratisSmith et al., 2016
    • AlopecocyonCamp and Vanderhoof, 1940
      • Alopecocyon leptorhynchus(Filhol, 1881) [syn.Alopecocyon goeriachensis]
    • ProtursusCrusafont & Kurtén, 1976
      • Protursus simpsoniCrusafont & Kurtén, 1976
    • SimocyonWagner, 1858
      • Simocyon batalleriViret, 1929
      • Simocyon diaphorus(Kaup, 1832)
      • Simocyon hungaricusKadic & Kretzoi, 1927
      • Simocyon primigenius(Roth & Wagner, 1854)
      • Simocyon marshiThorpe, 1921
  • Subfamily Ailurinae Gray, 1843
    • †ParailurusSchlosser, 1899
      • Parailurus anglicus(Dawkins, 1888)
      • Parailurus tedfordiWallace & Lyon, 2022
      • Parailurus baikalicusSotnikova, 2008
    • †PristinailurusWallace & Wang, 2004
      • Pristinailurus bristoliWallace & Wang, 2004
    • AilurusCuvier, 1825
      • Ailurus fulgens, the modern red panda

Taxonomic history

The species Ailurus fulgens was described by George Cuvier in 1825. ^[20]

In 1843, J.E. Gray assigned Ailurus fulgens to the new subfamily Ailurinae. ^[21]

In 1853, Pomel described the new genus Amphictis for "Viverra" antiqua. ^[22]

In 1858 the genus Simocyon was described. ^[23]

The subfamily Simocyoninae was named in 1868 for the genus Simocyon ^[24]

In 1899 the genus Parailurus was named. ^[25]

In 1940 the new genus Alopecocyon was described. ^[26]

In 1947 the new genus Actiocyon was described. ^[27]

In 1976 the new genus Protursus was described. ^[28]

In 1997 the new genus Magerictis was described. ^[29]

An additional, unnamed taxon called only "Ailurinae indet." was described in 2001 based on an upper molar from Four, a Middle Miocene-age locality near Isère, France. ^[30]

In 2004 the genus Pristinailurus was named. ^[15]

The new genera Bonisictis and Rothictis were described in 2025, both for species previously assigned to Amphictis. That genus along with Bonictis were assigned to the ressurected family Amphictidae, while Rothictis and Magerictis were assigned to the new ailurid subfamily Magerictinae. ^[19]

References

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4. Slattery, J.P.; O'Brien, S.J. (1995). "Molecular Phylogeny of the Red Panda (Ailurus fulgens)". J. Hered. 86 (6): 413–422. doi:10.1093/oxfordjournals.jhered.a111615. PMID   8568209.
5. Bing, Su; Fu, Yunxin; Wang, Yingxiang; Jin, Li; Chakraborty, Ranajit (2001). "Genetic Diversity and Population History of the Red Panda (Ailurus fulgens) as Inferred from Mitochondrial DNA Sequence Variations". Molecular Biology and Evolution. 18 (6): 1070–1076. doi: 10.1093/oxfordjournals.molbev.a003878 . PMID   11371595.
6. Li, Wen-Hsiung (2007). Molecular Evolution. Sinauer Associates. ISBN   978-0-87893-480-5.^{[ page needed ]}
7. Flynn et al., 2001^{[ full citation needed ]}
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9. Salesa, Manuel J.; Peigné, Stéphane; Antón, Mauricio; Morales, Jorge (2022). "Evolution of the family Ailuridae: origins and Old-World fossil record". Red Panda: Biology and Conservation of the First Panda (2nd ed.). Elsevier. pp. 15–29. doi:10.1016/b978-0-12-823753-3.00007-7. ISBN   978-0-12-823753-3. S2CID   243825496.
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11. Law, Chris J.; Slater, Graham J.; Mehta, Rita S. (2018-01-01). "Lineage Diversity and Size Disparity in Musteloidea: Testing Patterns of Adaptive Radiation Using Molecular and Fossil-Based Methods". Systematic Biology. 67 (1): 127–144. doi: 10.1093/sysbio/syx047 . ISSN   1063-5157. PMID   28472434.
12. McKenna, M.C.; Bell, S.K. (1997). Classification of Mammals Above the Species Level. Columbia University Press. ISBN   978-0-231-52853-5.^{[ page needed ]}
13. Peigné, S.; Salesa, M.; Antón, M.; Morales, J. (2005). "Ailurid carnivoran mammal Simocyon from the late Miocene of Spain and the systematics of the genus" (PDF). Acta Palaeontologica Polonica. 50: 219–238.
14. Salesa, M.; Antón, M.; Peigné, S.; Morales, J. (2006). "Evidence of a false thumb in a fossil carnivore clarifies the evolution of pandas". Proceedings of the National Academy of Sciences. 103 (2): 379–382. Bibcode:2006PNAS..103..379S. doi: 10.1073/pnas.0504899102 . PMC   1326154 . PMID   16387860.
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17. Baskin, Jon A. (2017). "Additional carnivorans from the early Hemingfordian Miller Local Fauna, Florida". Journal of Vertebrate Paleontology. 37 (2) e1293069. Bibcode:2017JVPal..37E3069B. doi:10.1080/02724634.2017.1293069. S2CID   90182619.
18. Smith, Kent; Czaplewski, Nicholas; Cifelli, Richard (2016). "Middle Miocene carnivorans from the Monarch Mill Formation, Nevada". Acta Palaeontologica Polonica. 61 (1): 231–252. Bibcode:2016AcPaP..61..231S. doi: 10.4202/app.00111.2014 .
19. Morales, Jorge; Abella, Juan; Caballero, Oscar; Demiguel, Daniel; Peláez-Campomanes, Pablo; Valenciano, Alberto (2025). "The enigmatic ailurid Magerictis imperialensis (Mammalia: Carnivora) unveiled: A systematic approach to the early Ailuridae". Journal of Systematic Palaeontology. 23 (1) 2571254. Bibcode:2025JSPal..2371254M. doi:10.1080/14772019.2025.2571254.
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Further reading

• Davis, D. Dwight (1964). The Giant Panda: A Morphological Study of Evolutionary Mechanisms. Fieldiana: Zoology Memoirs. Vol. 3. Chicago Natural History Museum Press. LCCN   64-8995. OCLC   1895716.
• Decker, D. M.; Wozencraft, W. C. (1991). "Phylogenetic Analysis of Recent Procyonid Genera". Journal of Mammalogy. 72 (1): 42–55. doi:10.2307/1381979. JSTOR   1381979.
• Flynn, J. J.; Hunt, G. D. Wesley (2005). "Carnivora". In Archibold, D.; Rose, K. (eds.). The Rise of Placental Mammals: Origin, Timing and Relationships of the Major Extant Clades. Baltimore: JHU Press. ISBN   0-8018-8022-X.
• Flynn, John J.; Finarelli, John A.; Zehr, Sarah; Hsu, Johnny; Nedbal, Michael A. (2005). "Molecular phylogeny of the Carnivora (Mammalia): ASS-ASS the impact of increased sampling to on resolving enigmatic relationships". Systematic Biology. 54 (2): 1–21. doi: 10.1080/10635150590923326 . PMID   16012099.
• Flynn, John J.; Nedbal, Michael A.; Dragoo, J. W.; Honeycutt, R. L. (November 2000). "Whence the Red Panda?" (PDF). Molecular Phylogenetics and Evolution. 17 (2): 190–199. Bibcode:2000MolPE..17..190F. doi:10.1006/mpev.2000.0819. PMID   11083933. Archived from the original (PDF) on 2006-09-10.
• Glatston, A. R., ed. (1989). Red Panda Biology. The Hague: SPB Academic Publishing. ISBN   90-5103-026-6.
• Glatston, A. R. (1994). The Red Panda, Olingos, Coatis, Raccoons, and their Relatives: Status survey and conservation action plan for Procyonids and Ailurids (PDF). IUCN/SSC Action Plans for the Conservation of Biological Diversity. Gland, Switzerland: IUCN. ISBN   978-2-8317-0046-5.
• Gregory, W. K. (August 8, 1936). "On the Phylogenetic Relationships of the Giant Panda (Ailuropoda) to other Arctoid Carnivores". American Museum Novitates (878): 1–29. hdl:2246/4138.
• Hu, J. C. (1990). 關於小熊貓的研究論文集[Proceedings of studies of the red panda] (in Chinese). Beijing: Chinese Scientific Publishing.
• Wilson, Don E.; Reeder, DeeAnn M. (2005). Mammal Species of the World: A Taxonomic and Geographic Reference. Vol. 1. JHU Press. ISBN   0-8018-8221-4.