Ailuridae: Difference between revisions

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
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Carnivora
Superfamily:	Musteloidea
Family:	Ailuridae; Gray, 1843
Subfamilies
	†Amphictinae?; †Simocyoninae; Ailurinae;
	Extant red panda distribution

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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) and a group that further subdivides into 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]

Caniformia

Arctoidea

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 borbonica Viret, 1929
    - †Amphictis ambigua (Gervais, 1872)
    - †Amphictis milloquensis (Helbing, 1936)
    - †Amphictis antiqua (de Blainville, 1842)
    - †Amphictis schlosseri Heizmann & Morlo, 1994
    - †Amphictis prolongata Morlo, 1996
    - †Amphictis wintershofensis Roth, 1994
    - †Amphictis cuspida Nagel, 2003
    - †Amphictis timucua J.A. Baskin, 2017^[17]
- Subfamily †Simocyoninae Dawkins, 1868
  - †Actiocyon Stock, 1947
    - †Actiocyon parverratis Smith et al., 2016^[18]
    - †Actiocyon leardi Stock, 1947
  - †Alopecocyon Camp & Vanderhoof, 1940
    - †Alopecocyon getti Mein, 1958
    - †Alopecocyon goeriachensis (Toula, 1884)
  - †Protursus Crusafont & Kurtén, 1976
    - †Protursus simpsoni Crusafont & Kurtén, 1976
  - †Simocyon Wagner, 1858
    - †Simocyon primigenius (Roth & Wagner, 1854)
    - †Simocyon diaphorus (Kaup, 1832)
    - †Simocyon batalleri Viret, 1929
    - †Simocyon hungaricus Kadic & Kretzoi, 1927
- Subfamily Ailurinae J.E. Gray, 1843
  - †Magerictis Ginsburg et al., 1997
    - †Magerictis imperialensis Ginsburg et al., 1997
  - Tribe Pristinailurini Wallace & Lyon, 2022
    - †Pristinailurus Wallace & Wang, 2004
      - †Pristinailurus bristoli Wallace & Wang, 2004
    - †Parailurus Schlosser, 1899
      - †Parailurus anglicus (Dawkins, 1888) [Parailurus hungaricus Kormos, 1935]
      - †Parailurus tedfordi Wallace & Lyon, 2022
      - †Parailurus baikalicus Sotnikova, 2008
  - Tribe Ailurini
    - Ailurus F. Cuvier, 1825
      - Ailurus fulgens - Red panda
        Ailurus fulgens styani Thomas, 1902 – Eastern red panda
        
        Ailurus fulgens fulgens F. Cuvier, 1825 – Western red panda

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.^[19]

References

^ ^a ^b ^c Mayr, E. (1986). "Uncertainty in Science: is the Giant panda a bear or a raccoon?". Nature. 323 (6091): 769–771. Bibcode:1986Natur.323..769M. doi:10.1038/323769a0. PMID 3774006. S2CID 4317414.
^ Roberts, M.S.; Gittleman, J.L. (1984). "Ailurus fulgens". Mammalian Species (222). American Society of Mammalogists: 1–8. doi:10.2307/3503840. JSTOR 3503840.
^ Zhang, Y.P.; Ryder, O.A. (1993). "Mitochondrial DNA sequence evolution in the Arctoidea". PNAS. 90 (20): 9557–9561. Bibcode:1993PNAS...90.9557Z. doi:10.1073/pnas.90.20.9557. PMC 47608. PMID 8415740.
^ 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.
^ 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.
^ Li, Wen-Hsiung (2007). Molecular Evolution. Sinauer Associates. ISBN 978-0-87893-480-5.^{[page needed]}
^ Flynn et al., 2001^{[full citation needed]}
^ Flynn, John J.; Nedbal, Michael A.; Dragoo, Jerry W.; Honeycutt, Rodney L. (1 November 2000). "Whence the Red Panda?" (PDF). Molecular Phylogenetics and Evolution. 17 (2): 190–99. doi:10.1006/mpev.2000.0819. ISSN 1055-7903. PMID 11083933. Archived from the original (PDF) on 13 April 2018. Retrieved 11 February 2018.
^ Salesa, Manuel J.; Peigné, Stéphane; Antón, Mauricio; Morales, Jorge (2022), "Evolution of the family Ailuridae: origins and Old-World fossil record", Red Panda, Elsevier, pp. 15–29, doi:10.1016/b978-0-12-823753-3.00007-7, ISBN 9780128237533, S2CID 243825496, retrieved 2022-06-29
^ Flynn, J.J.; Finarelli, J.A.; Zehr, S.; Hsu, J.; Nedbal, M.A. (2005). "Molecular phylogeny of the Carnivora (Mammalia): Assessing the impact of increased sampling on resolving enigmatic relationships". Systematic Biology. 54 (2): 317–337. doi:10.1080/10635150590923326. PMID 16012099.
^ 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.
^ McKenna, M.C.; Bell, S.K. (1997). Classification of Mammals Above the Species Level. Columbia University Press. ISBN 9780231528535.^{[page needed]}
^ 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.
^ 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.
^ Wallace, S.C.; Wang, X. (2004). "Two new carnivores from an unusual late Tertiary forest biota in eastern North America". Nature. 431 (7008): 556–559. Bibcode:2004Natur.431..556W. doi:10.1038/nature02819. PMID 15457257. S2CID 4432191.
^ Morlo, Michael; Peigné, Stéphane (2010). "Molecular and morphological evidence for Ailuridae and a review of its genera". In Goswami, Anjali; Friscia, Anthony (eds.). Carnivoran Evolution: New Views on Phylogeny, Form, and Function. pp. 92–140. doi:10.1017/CBO9781139193436.005. ISBN 978-0-521-73586-5.
^ 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.
^ Smith, Kent; Czaplewski, Nicholas; Cifelli, Richard (2016). "Middle Miocene carnivorans from the Monarch Mill Formation, Nevada". Acta Palaeontologica Polonica. 61 (1): 231–252. doi:10.4202/app.00111.2014.
^ Ginsburg, Leonard; Maridet, Olivier; Mein, Pierre (2001). "Un Ailurinae (Mammalia, Carnivora, Ailuridae) dans le Miocène moyen de Four (Isère, France)" [An Ailurinae (Mammalia, Carnivora, Ailuridae) in the middle Miocene of Four (Isère, France)]. Geodiversitas (in French). 23 (1): 81–85.

@@ Line 4: / Line 4: @@
 | image = Ailurus fulgens RoterPanda LesserPanda.jpg
 | image_caption = [[Red panda]] (''Ailurus fulgens'')
-| image2 = Skull of Pristinailurus bristoli 2.jpg
+| image2 = Simocyon batalleri head.jpg
-| image2_caption = Skull of ''[[Pristinailurus bristoli]]''
+| image2_caption = Skull and life restoration of ''[[Simocyon]]''
 | taxon = Ailuridae
 | authority = [[John Edward Gray|Gray]], 1843
@@ Line 18: / Line 18: @@
 '''Ailuridae''' is a [[family (biology)|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 [[Procyonidae|raccoon family]] in 1825; this classification has been controversial ever since.<ref name=Mayr>{{cite journal |author-last=Mayr |author-first=E. |year=1986 |title=Uncertainty in Science: is the Giant panda a bear or a raccoon? |journal=Nature |volume=323 |pages=769–771 |doi=10.1038/323769a0 |pmid=3774006 |issue=6091|bibcode=1986Natur.323..769M |s2cid=4317414 }}</ref> It was classified in the raccoon family because of [[morphology (biology)|morphological]] similarities of the head, colored ringed tail, and other morphological and [[ecological]] characteristics. Somewhat later, it was assigned to the [[bear]] family.<ref name=Roberts/>
+[[Georges Cuvier]] first described ''Ailurus'' as belonging to the [[Procyonidae|raccoon family]] in 1825; this classification has been controversial ever since.<ref name=Mayr>{{cite journal |author-last=Mayr |author-first=E. |year=1986 |title=Uncertainty in Science: is the Giant panda a bear or a raccoon? |journal=Nature |volume=323 |pages=769–771 |doi=10.1038/323769a0 |pmid=3774006 |issue=6091|bibcode=1986Natur.323..769M |s2cid=4317414 }}</ref> It was classified in the raccoon family because of [[morphology (biology)|morphological]] similarities of the head, colored ringed tail, and other morphological and [[ecological]] characteristics. Somewhat later, it was assigned to the [[bear]] family.<ref name="Roberts">{{cite journal |year=1984 |title=''Ailurus fulgens'' |journal=Mammalian Species |publisher=American Society of Mammalogists |issue=222 |pages=1–8 |doi=10.2307/3503840 |jstor=3503840 |author-last1=Roberts |author-first1=M.S. |author-last2=Gittleman |author-first2=J.L.|doi-access=free }}</ref>
 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.<ref name=Mayr/><ref name="Zhang">{{cite journal |author-last1=Zhang |author-first1=Y.P. |author-last2=Ryder |author-first2=O.A. |year=1993 |title=Mitochondrial DNA sequence evolution in the Arctoidea |journal=[[PNAS]] |volume=90 |issue=20 |pages=9557–9561 |doi=10.1073/pnas.90.20.9557 |pmc=47608 |pmid=8415740|bibcode=1993PNAS...90.9557Z |doi-access=free }}</ref><ref name="Slattery">{{cite journal |author-last1=Slattery |author-first1=J.P. |author-last2=O'Brien |author-first2=S.J. |year=1995 |title=Molecular Phylogeny of the Red Panda (''Ailurus fulgens'') |journal=J. Hered. |volume=86 |issue=6 |pages=413–422 |pmid=8568209 |doi=10.1093/oxfordjournals.jhered.a111615}}</ref> 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."<ref name="Mayr" /><ref name=Bing>{{cite journal |author-first1=Su |author-last1=Bing |author-first2=Yunxin |author-last2=Fu |author-first3=Yingxiang |author-last3=Wang |author-first4=Li |author-last4=Jin |author-first5=Ranajit |author-last5=Chakraborty |year=2001 |title=Genetic Diversity and Population History of the Red Panda (Ailurus fulgens) as Inferred from Mitochondrial DNA Sequence Variations |journal=Molecular Biology and Evolution |volume=18 |pages=1070–1076 |pmid=11371595 |issue=6 |doi=10.1093/oxfordjournals.molbev.a003878|doi-access=free }}</ref> With this divergence, by comparing the [[DNA sequencing|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 10<sup>9</sup>, which is apparently an underestimate compared with the average rate in mammals.<ref>{{cite book |title=Molecular Evolution |author-last=Li |author-first=Wen-Hsiung |publisher=Sinauer Associates |year=2007 |isbn=978-0-87893-480-5}}{{Page needed |date=February 2018}}</ref> This underestimation is probably due to multiple recurrent mutations as the [[divergence]] between the red panda and the raccoon is extremely deep.{{Citation needed|date=May 2019}}
@@ Line 24: / Line 24: @@
 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]]<ref>Flynn et al., 2001{{Full citation needed|date=February 2018}}</ref> that also includes the [[Procyonidae]] ([[raccoon]]s) and a group that further subdivides into the Mephitidae ([[skunk]]s) and [[Mustelidae]] ([[weasel]]s); but it is not a [[bear]] (Ursidae).<ref>{{cite journal |author-last1=Flynn |author-first1=John J. |author-last2=Nedbal |author-first2=Michael A. |author-last3=Dragoo |author-first3=Jerry W. |author-last4=Honeycutt |author-first4=Rodney L. |date=1 November 2000 |title=Whence the Red Panda? |url=http://pandafriends.ru/wp-content/uploads/2011/01/Flynn2000.pdf |journal=Molecular Phylogenetics and Evolution |volume=17 |issue=2 |pages=190–99 |doi=10.1006/mpev.2000.0819 |issn=1055-7903 |access-date=11 February 2018 |archive-url=https://web.archive.org/web/20180413005734/http://pandafriends.ru/wp-content/uploads/2011/01/Flynn2000.pdf |archive-date=13 April 2018 |url-status=dead |pmid=11083933}}</ref>
+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 [[Cusp (anatomy)|cusps]], creating an effective grinding surface to process plant material.<ref>{{Citation |last1=Salesa |first1=Manuel J. |title=Evolution of the family Ailuridae: origins and Old-World fossil record |date=2022 |url=http://dx.doi.org/10.1016/b978-0-12-823753-3.00007-7 |work=Red Panda |pages=15–29 |publisher=Elsevier |access-date=2022-06-29 |last2=Peigné |first2=Stéphane |last3=Antón |first3=Mauricio |last4=Morales |first4=Jorge|doi=10.1016/b978-0-12-823753-3.00007-7 |isbn=9780128237533 |s2cid=243825496 }}</ref>
-Red pandas have no close living relatives, and their nearest fossil ancestors, ''Parailurus'', lived 3-4 million years ago. There may have been as many as three different species of ''Parailurus'', all larger and more [[Robustness (morphology)|robust]] in the head and jaw than ''Ailurus'', living in [[Eurasia]] and possibly crossing the [[Bering Strait]] into the [[Americas]]. The red panda may be the sole surviving species – a specialized offshoot surviving the [[last glacial period]] in a Chinese mountain refuge.<ref name=Roberts>{{cite journal |author-last1=Roberts |author-first1=M.S. |author-last2=Gittleman |author-first2=J.L. |year=1984 |title=''Ailurus fulgens'' |journal=Mammalian Species |volume=222 |issue=222 |pages=1–8 |doi=10.2307/3503840 |jstor=3503840 |publisher=American Society of Mammalogists}}</ref> The thick coat of fur allows for the red panda to survive harsh environmental changes and cold temperatures.<ref>{{Cite web|url=https://nationalzoo.si.edu/animals/red-panda|title = Red panda|date = 22 April 2016}}</ref>
 ==Classification==
+The relationship of the Ailuridae with other carnivorans is shown in the following [[cladogram|phylogenetic tree]], which is based on the [[molecular phylogeny|molecular phylogenetic analysis]] of six genes in Flynn (2005),<ref name="Flynn2005">{{Cite journal |author1=Flynn, J.J. |author2=Finarelli, J.A. |author3=Zehr, S. |author4=Hsu, J. |author5=Nedbal, M.A. |title=Molecular phylogeny of the Carnivora (Mammalia): Assessing the impact of increased sampling on resolving enigmatic relationships |doi=10.1080/10635150590923326 |journal=Systematic Biology |volume=54 |issue=2 |pages=317–337 |year=2005 |pmid=16012099|doi-access=free }}</ref> with the musteloids updated following the multigene analysis of Law et al. (2018).<ref name="Law-2018">{{Cite journal|last=Law|first=Chris J.|last2=Slater|first2=Graham J.|last3=Mehta|first3=Rita S.|date=2018-01-01|title=Lineage Diversity and Size Disparity in Musteloidea: Testing Patterns of Adaptive Radiation Using Molecular and Fossil-Based Methods|journal=Systematic Biology|language=en|volume=67|issue=1|pages=127–144|doi=10.1093/sysbio/syx047|pmid=28472434|issn=1063-5157|doi-access=free}}</ref>
-In addition to ''Ailurus'', the family Ailuridae includes seven extinct genera, most of which are assigned to three subfamilies: Amphicinae, Simocyoninae, and Ailurinae.<ref name="McKenna">{{cite book|title=Classification of Mammals Above the Species Level |author-last1=McKenna |author-first1=M.C. |author-last2=Bell |author-first2=S.K. |publisher=Columbia University Press |year=1997 |isbn=9780231528535}}{{Page needed|date=February 2018}}</ref><ref name="Peigne">{{cite journal |author-last1=Peigné |author-first1=S. |author-last2=Salesa |author-first2=M. |author-last3=Antón |author-first3=M. |author-last4=Morales |author-first4=J. |year=2005 |title=Ailurid carnivoran mammal ''Simocyon'' from the late Miocene of Spain and the systematics of the genus |url=http://agro.icm.edu.pl/agro/element/bwmeta1.element.agro-article-8012e086-056a-483a-821e-01d84489ff00/c/app50-219.pdf |journal=Acta Palaeontologica Polonica |volume=50 |pages=219–238}}</ref><ref name=Salesa>{{cite journal |author-last1=Salesa |author-first1=M. |author-last2=Antón |author-first2=M. |author-last3=Peigné |author-first3=S. |author-last4=Morales |author-first4=J. |year=2006 |title=Evidence of a false thumb in a fossil carnivore clarifies the evolution of pandas |journal=Proceedings of the National Academy of Sciences |volume=103 |pages=379–382 |doi=10.1073/pnas.0504899102 |pmid=16387860 |issue=2 |pmc=1326154|bibcode=2006PNAS..103..379S |doi-access=free }}</ref><ref name="Wallace">{{cite journal |author-last1=Wallace |author-first1=S.C. |author-last2=Wang |author-first2=X. |year=2004 |title=Two new carnivores from an unusual late Tertiary forest biota in eastern North America |url=https://www.researchgate.net/publication/8258760 |journal=Nature |volume=431 |issue=7008 |pages=556–559 |doi=10.1038/nature02819 |pmid=15457257 |bibcode=2004Natur.431..556W |s2cid=4432191}}</ref><ref>{{cite book |title=Carnivoran Evolution: New Views on Phylogeny, Form, and Function |author-last1=Morlo |author-first1=Michael |author-last2=Peigné |author-first2=Stéphane |editor1-first=Anjali |editor1-last=Goswami |editor2-first=Anthony |editor2-last=Friscia |year=2010 |isbn=978-0-521-73586-5 |pages=92–140 |chapter=Molecular and morphological evidence for Ailuridae and a review of its genera |doi=10.1017/CBO9781139193436.005}}</ref>
+{{clade|{{clade
+   |1=[[Feliformia]] [[File:Lydekker - Ocelot (white background).JPG|60px|Ocelot]]
+   |label2=&nbsp;&nbsp;&nbsp;[[Caniformia]]&nbsp;&nbsp;&nbsp;
+   |2={{clade
+      |1=[[Canidae]] [[File:Dogs, jackals, wolves, and foxes (Plate XI).jpg|60px|African golden wolf]]
+      |label2=&nbsp;&nbsp;&nbsp;[[Arctoidea]]&nbsp;&nbsp;&nbsp;
+      |2={{clade
+         |label1=&nbsp;&nbsp;&nbsp;
+         |1=[[Ursidae]] [[File:Ursus arctos - 1700-1880 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - (white background).jpg|70px|Brown bear]]
+         |2={{clade
+            |1=[[Pinnipedia]]  [[File:Faroe stamp 227 grey seal (Phoca vitulina) white background.jpg|60px|Common seal]]
+            |label2=[[Musteloidea]]
+            |2={{clade
+               |1=[[Mephitidae]] [[File:Die Säugthiere in Abbildungen nach der Natur, mit Beschreibungen (Plate CXXI-) (white background).jpg|50px|Striped skunk]]
+               |2='''Ailuridae''' [[File:Ailurus fulgens - 1700-1880 - Print - Iconographia Zoologica - Special Collections University of Amsterdam -(white background).jpg|60px|Red panda]]
+               |3={{clade
+                  |1=[[Procyonidae]] [[File:Wild animals of North America, intimate studies of big and little creatures of the mammal kingdom (Page 410) (white background).jpg|60 px|Common raccoon]]
+                  |2=[[Mustelidae]] [[File:Mustela eversmanii (white background).png|60px|Steppe polecat]]
+                  }}
+               }}
+            }}
+         }}
+      }}
+   }}|label1=[[Carnivora]]}}
+In addition to ''Ailurus'', the family Ailuridae includes seven extinct genera, most of which are assigned to three subfamilies: Amphictinae, Simocyoninae, and Ailurinae.<ref name="McKenna">{{cite book|title=Classification of Mammals Above the Species Level |author-last1=McKenna |author-first1=M.C. |author-last2=Bell |author-first2=S.K. |publisher=Columbia University Press |year=1997 |isbn=9780231528535}}{{Page needed|date=February 2018}}</ref><ref name="Peigne">{{cite journal |author-last1=Peigné |author-first1=S. |author-last2=Salesa |author-first2=M. |author-last3=Antón |author-first3=M. |author-last4=Morales |author-first4=J. |year=2005 |title=Ailurid carnivoran mammal ''Simocyon'' from the late Miocene of Spain and the systematics of the genus |url=http://agro.icm.edu.pl/agro/element/bwmeta1.element.agro-article-8012e086-056a-483a-821e-01d84489ff00/c/app50-219.pdf |journal=Acta Palaeontologica Polonica |volume=50 |pages=219–238}}</ref><ref name=Salesa>{{cite journal |author-last1=Salesa |author-first1=M. |author-last2=Antón |author-first2=M. |author-last3=Peigné |author-first3=S. |author-last4=Morales |author-first4=J. |year=2006 |title=Evidence of a false thumb in a fossil carnivore clarifies the evolution of pandas |journal=Proceedings of the National Academy of Sciences |volume=103 |pages=379–382 |doi=10.1073/pnas.0504899102 |pmid=16387860 |issue=2 |pmc=1326154|bibcode=2006PNAS..103..379S |doi-access=free }}</ref><ref name="Wallace">{{cite journal |author-last1=Wallace |author-first1=S.C. |author-last2=Wang |author-first2=X. |year=2004 |title=Two new carnivores from an unusual late Tertiary forest biota in eastern North America |url=https://www.researchgate.net/publication/8258760 |journal=Nature |volume=431 |issue=7008 |pages=556–559 |doi=10.1038/nature02819 |pmid=15457257 |bibcode=2004Natur.431..556W |s2cid=4432191}}</ref><ref>{{cite book |title=Carnivoran Evolution: New Views on Phylogeny, Form, and Function |author-last1=Morlo |author-first1=Michael |author-last2=Peigné |author-first2=Stéphane |editor1-first=Anjali |editor1-last=Goswami |editor2-first=Anthony |editor2-last=Friscia |year=2010 |isbn=978-0-521-73586-5 |pages=92–140 |chapter=Molecular and morphological evidence for Ailuridae and a review of its genera |doi=10.1017/CBO9781139193436.005}}</ref>
-* Family Ailuridae <small>J.E. Gray, 1856</small>
+* Family Ailuridae <small>J.E. Gray, 1843</small>
 ** Subfamily †[[Amphictis|Amphictinae]] ?<small>Winge, 1896</small>
 *** †''[[Amphictis]]'' ?<small>Pomel, 1853</small>
@@ Line 40: / Line 66: @@
 **** †''Amphictis wintershofensis'' <small>Roth, 1994</small>
 **** †''Amphictis cuspida'' <small>Nagel, 2003</small>
-**** †''Amphictis timucua'' <small>J.A. Baskin, 2017</small><ref name=Floridictis>{{cite journal|author-first=Jon A. |author-last=Baskin |year=2017 |title=Additional carnivorans from the early Hemingfordian Miller Local Fauna, Florida |journal=Journal of Vertebrate Paleontology |volume=37 |issue=2 |pages=e1293069 |doi=10.1080/02724634.2017.1293069 |s2cid=90182619 }}</ref>
+**** †''Amphictis timucua'' <small>J.A. Baskin, 2017</small><ref name=Floridictis>{{cite journal|author-first=Jon A. |author-last=Baskin |year=2017 |title=Additional carnivorans from the early Hemingfordian Miller Local Fauna, Florida |journal=Journal of Vertebrate Paleontology |volume=37 |issue=2 |pages=e1293069 |doi=10.1080/02724634.2017.1293069 |bibcode=2017JVPal..37E3069B |s2cid=90182619 }}</ref>
 ** Subfamily †[[Simocyoninae]] <small>Dawkins, 1868</small>
 *** †''[[Actiocyon]]'' <small>Stock, 1947</small>
@@ Line 48: / Line 74: @@
 **** †''Alopecocyon getti'' <small>Mein, 1958</small>
 **** †''Alopecocyon goeriachensis'' <small>(Toula, 1884)</small>
-*** †''Protursus'' <small>Crusafont & Kurtén, 1976</small>
+*** †''[[Protursus]]'' <small>Crusafont & Kurtén, 1976</small>
 **** †''Protursus simpsoni'' <small>Crusafont & Kurtén, 1976</small>
 *** †''[[Simocyon]]'' <small>Wagner, 1858</small>
@@ Line 56: / Line 82: @@
 **** †''Simocyon hungaricus'' <small>Kadic & Kretzoi, 1927</small>
 ** Subfamily [[Ailurinae]] <small>J.E. Gray, 1843</small>
-*** †''Magerictis'' <small>Ginsburg et al., 1997</small>
+*** †''[[Magerictis]]'' <small>Ginsburg et al., 1997</small>
 **** †''Magerictis imperialensis'' <small>Ginsburg et al., 1997</small>
 *** Tribe Pristinailurini <small>Wallace & Lyon, 2022</small>
@@ Line 67: / Line 93: @@
 *** Tribe Ailurini
 **** ''[[Ailurus]]'' <small>F. Cuvier, 1825</small>
-***** ''Ailurus fulgens'' - Red Panda
+***** ''Ailurus fulgens'' - Red panda
-****** ''Ailurus fulgens styani'' <small>Thomas, 1902</small> – Eastern Red Panda
+****** ''Ailurus fulgens styani'' <small>Thomas, 1902</small> – Eastern red panda
-****** ''Ailurus fulgens fulgens'' <small>F. Cuvier, 1825</small> – Western Red Panda
+****** ''Ailurus fulgens fulgens'' <small>F. Cuvier, 1825</small> – Western red panda
+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]].<ref>{{cite journal |last1=Ginsburg |first1=Leonard |last2=Maridet |first2=Olivier |last3=Mein |first3=Pierre |date=2001 |title=Un Ailurinae (Mammalia, Carnivora, Ailuridae) dans le Miocène moyen de Four (Isère, France) |trans-title=An Ailurinae (Mammalia, Carnivora, Ailuridae) in the middle Miocene of Four (Isère, France) |url=https://www.researchgate.net/publication/237599256 |language=French |journal=Geodiversitas |volume=23 |issue=1 |pages=81–85 |doi= |access-date=}}</ref>
 ==References==
@@ Line 75: / Line 103: @@
 ==Further reading==
-* {{cite journal | author = Davis Davis D | year = 1964 | title = The Giant Panda: A Morphological Study of Evolutionary Mechanisms | journal = Zoology Memoirs | volume = 3 | pages = 1–339 }}
+* {{cite journal | author = Davis D. Dwight | year = 1964 | title = The Giant Panda: A Morphological Study of Evolutionary Mechanisms | journal = Zoology Memoirs | volume = 3 | pages = 1–339 }}
 * {{cite journal | author = Decker D.M., Wozencraft W.C. | year = 1991 | title = Phylogenetic Analysis of Recent Procyonid Genera | journal = Journal of Mammalogy | volume = 72 | issue = 1| pages = 42–55 | doi=10.2307/1381979| jstor = 1381979 }}
 * Flynn, J.J. and G.D. Wesley Hunt. (2005a). "Carnivora." in ''The Rise of Placental Mammals: Origin, Timing and Relationships of the Major Extant Clades'', by D. Archibold and K. Rose. Baltimore. {{ISBN|0-8018-8022-X}}
@@ Line 94: / Line 122: @@
 [[Category:Carnivorans]]
 [[Category:Mammal families]]
-[[Category:Oligocene first appearances]]
+[[Category:Extant Oligocene first appearances]]
 [[Category:Taxa named by John Edward Gray]]