Ailurid
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Ailuridae is a
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
in the
mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
order
Carnivora Carnivora ( ) is an order of placental mammals specialized primarily in eating flesh, whose members are formally referred to as carnivorans. The order Carnivora is the sixth largest order of mammals, comprising at least 279 species. Carnivor ...
. The family consists of the
red panda The red panda (''Ailurus fulgens''), also known as the lesser panda, is a small mammal native to the eastern Himalayas and southwestern China. It has dense reddish-brown fur with a black belly and legs, white-lined ears, a mostly white muzz ...
(the sole living representative) and its extinct relatives.
Georges Cuvier Jean Léopold Nicolas Frédéric, baron Cuvier (23 August 1769 – 13 May 1832), known as Georges Cuvier (; ), was a French natural history, naturalist and zoology, zoologist, sometimes referred to as the "founding father of paleontology". Cuv ...
first described ''Ailurus'' as belonging to the raccoon family in 1825; this classification has been controversial ever since. It was classified in the raccoon family because of morphological similarities of the head, colored ringed tail, and other morphological and
ecological Ecology () is the natural science of the relationships among living organisms and their environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere levels. Ecology overlaps with the closely re ...
characteristics. Somewhat later, it was assigned to the
bear Bears are carnivoran mammals of the family (biology), family Ursidae (). They are classified as caniforms, or doglike carnivorans. Although only eight species of bears are extant, they are widespread, appearing in a wide variety of habitats ...
family. Molecular
phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
studies had shown that, as an ancient species in the order
Carnivora Carnivora ( ) is an order of placental mammals specialized primarily in eating flesh, whose members are formally referred to as carnivorans. The order Carnivora is the sixth largest order of mammals, comprising at least 279 species. Carnivor ...
, the red panda is relatively close to the American
raccoon The raccoon ( or , ''Procyon lotor''), sometimes called the North American, northern or common raccoon (also spelled racoon) to distinguish it from Procyonina, other species of raccoon, is a mammal native to North America. It is the largest ...
and may be either a monotypic family or a subfamily within the
procyonid Procyonidae ( ) is a New World family (biology), family of the order (biology), order Carnivora. It includes the Procyon (genus), raccoons, ringtails, cacomistles, coatis, kinkajous, olingos, and olinguitos. Procyonids inhabit a wide range of e ...
family. An in-depth
mitochondrial DNA Mitochondrial DNA (mtDNA and mDNA) is the DNA located in the mitochondrion, mitochondria organelles in a eukaryotic cell that converts chemical energy from food into adenosine triphosphate (ATP). Mitochondrial DNA is a small portion of the D ...
population analysis study stated: "According to the fossil record, the Red Panda diverged from its common ancestor with bears about 40 million years ago." With this divergence, by comparing the
sequence In mathematics, a sequence is an enumerated collection of objects in which repetitions are allowed and order matters. Like a set, it contains members (also called ''elements'', or ''terms''). The number of elements (possibly infinite) is cal ...
difference between the red panda and the raccoon, the observed
mutation rate In genetics, the mutation rate is the frequency of new mutations in a single gene, nucleotide sequence, or organism over time. Mutation rates are not constant and are not limited to a single type of mutation; there are many different types of mu ...
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. This underestimation is probably due to multiple recurrent mutations as the
divergence In vector calculus, divergence is a vector operator that operates on a vector field, producing a scalar field giving the rate that the vector field alters the volume in an infinitesimal neighborhood of each point. (In 2D this "volume" refers to ...
between the red panda and the raccoon is extremely deep. The most recent molecular-systematic
DNA Deoxyribonucleic acid (; DNA) is a polymer composed of two polynucleotide chains that coil around each other to form a double helix. The polymer carries genetic instructions for the development, functioning, growth and reproduction of al ...
research places the red panda into its own independent family, Ailuridae. Ailuridae are, in turn, part of a trichotomy within the broad superfamily
Musteloidea Musteloidea is a superfamily (taxonomy), superfamily of carnivoran mammals united by shared characteristics of the skull and teeth. Musteloids are the sister group of pinnipeds, the group which includes seals. Musteloidea comprises the following ...
that also includes the
Procyonidae Procyonidae ( ) is a New World family of the order Carnivora. It includes the raccoons, ringtails, cacomistles, coatis, kinkajous, olingos, and olinguitos. Procyonids inhabit a wide range of environments and are generally omnivorous. Ch ...
(
raccoon The raccoon ( or , ''Procyon lotor''), sometimes called the North American, northern or common raccoon (also spelled racoon) to distinguish it from Procyonina, other species of raccoon, is a mammal native to North America. It is the largest ...
s), the Mephitidae (
skunk Skunks are mammals in the family Mephitidae. They are known for their ability to spray a liquid with a strong, unpleasant scent from their anal glands. Different species of skunk vary in appearance from black-and-white to brown, cream or gi ...
s), and
Mustelidae The Mustelidae (; from Latin , weasel) are a diverse family of carnivora, carnivoran mammals, including weasels, badgers, otters, polecats, martens, grisons, and wolverines. Otherwise known as mustelids (), they form the largest family in the s ...
(
weasel Weasels are mammals of the genus ''Mustela'' of the family Mustelidae. The genus ''Mustela'' includes the least weasels, polecats, stoats, ferrets, and European mink. Members of this genus are small, active predators, with long and slend ...
s); but it is not a
bear Bears are carnivoran mammals of the family (biology), family Ursidae (). They are classified as caniforms, or doglike carnivorans. Although only eight species of bears are extant, they are widespread, appearing in a wide variety of habitats ...
(Ursidae). Ailurids appear to have originated during the Late
Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...
to Early
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
in Europe. The earliest known member, ''
Amphictis ''Amphictis'' is an extinct genus of ailurid that existed from the Late Oligocene to the Middle Miocene with fossils found in Eurasia and North America with a total of nine described species. The interrelationships of the different species as we ...
'', was likely an unspecialised carnivore, based on its dentition. Ailurids subsequently dispersed into Asia and North America. The puma-sized ''
Simocyon ''Simocyon'' ("short-snouted dog") is a genus of extinct carnivoran mammal in the family Ailuridae. ''Simocyon'', which was about the size of a mountain lion, lived in the late Miocene and early Pliocene epochs, and has been found in Europe, Asi ...
'' found in Middle
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
-Early
Pliocene The Pliocene ( ; also Pleiocene) is the epoch (geology), epoch in the geologic time scale that extends from 5.33 to 2.58hypercarnivore A hypercarnivore is an animal that has a diet that is more than 70% meat, either via active predation or by scavenging. The remaining non-meat diet may consist of non-animal foods such as fungi, fruits or other plant material. Some extant exampl ...
. 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 ''Pristinailurus bristoli'' is a fossil species in the carnivoran family Ailuridae, well-represented in the Hemphillian deposits at the Gray Fossil Site The Gray Fossil Site is an Early Pliocene assemblage of fossils dating between 4.5 and 4.9 ...
'' and '' Parailurus'', developed a specialised dental morphology with blunted cusps, creating an effective grinding surface to process plant material.


Classification

The relationship of the Ailuridae with other carnivorans is shown in the following
phylogenetic tree A phylogenetic tree or phylogeny is a graphical representation which shows the evolutionary history between a set of species or taxa during a specific time.Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA. In ...
, which is based on the molecular phylogenetic analysis of six genes in Flynn (2005), with the musteloids updated following the multigene analysis of Law et al. (2018). In addition to ''Ailurus'', the family Ailuridae includes seven extinct genera, most of which are assigned to three subfamilies: Amphictinae, Simocyoninae, and Ailurinae. * Family Ailuridae J.E. Gray, 1843 ** Subfamily † Amphictinae ?Winge, 1896 *** †''
Amphictis ''Amphictis'' is an extinct genus of ailurid that existed from the Late Oligocene to the Middle Miocene with fossils found in Eurasia and North America with a total of nine described species. The interrelationships of the different species as we ...
'' ?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 ** Subfamily †
Simocyoninae Simocyoninae is an extinct subfamily of Ailuridae (of which the only recent member is the red panda). The taxonomic history of this group was complicated, as researchers placed various fossil caniform genera into the subfamily. In addition to ''S ...
Dawkins, 1868 *** †''
Actiocyon ''Actiocyon'' was an extinct genus of ailurid that existed from the Middle Miocene of North America. The type species, ''Actiocyon leardi'', was described in 1947 by the American paleontologist Chester Stock. Description The holotype consisted ...
'' Stock, 1947 **** †''Actiocyon parverratis'' Smith et al., 2016 **** †''Actiocyon leardi'' Stock, 1947 *** †''
Alopecocyon ''Alopecocyon'' is a fossil genus of ailurid belonging to the subfamily Simocyoninae. It has two species, ''Alopecocyon goeriachensis'' and ''Alopecocyon getti''. It is based on fragmentary fossils dating to the middle Miocene The Miocene ( ...
'' Camp & Vanderhoof, 1940 **** †''Alopecocyon getti'' Mein, 1958 **** †''Alopecocyon goeriachensis'' (Toula, 1884) *** †''
Protursus ''Protursus'' is a fossil genus of ailurid with a single species ''Protursus simpsoni'', known from a single left m2 tooth. Taxonomy & Evolution ''Protursus simpsoni'' was believed to be an ursid (bear) closely related to ''Ursavus'' by its orig ...
'' Crusafont & Kurtén, 1976 **** †''Protursus simpsoni'' Crusafont & Kurtén, 1976 *** †''
Simocyon ''Simocyon'' ("short-snouted dog") is a genus of extinct carnivoran mammal in the family Ailuridae. ''Simocyon'', which was about the size of a mountain lion, lived in the late Miocene and early Pliocene epochs, and has been found in Europe, Asi ...
'' Wagner, 1858 **** †''Simocyon primigenius'' (Roth & Wagner, 1854) **** †''Simocyon diaphorus'' (Kaup, 1832) **** †''Simocyon batalleri'' Viret, 1929 **** †''Simocyon hungaricus'' Kadic & Kretzoi, 1927 ** Subfamily
Ailurinae Ailurinae is a subfamily of Ailuridae (of which this is the only extant subfamily out of three). While it is represented by the extant genus ''Ailurus'', there were a handful of genera whose fossils have been found across the Holarctic region. Th ...
J.E. Gray, 1843 *** †'' Magerictis'' Ginsburg et al., 1997 **** †''Magerictis imperialensis'' Ginsburg et al., 1997 *** Tribe Pristinailurini Wallace & Lyon, 2022 **** †''
Pristinailurus ''Pristinailurus bristoli'' is a fossil species in the carnivoran family Ailuridae, well-represented in the Hemphillian deposits at the Gray Fossil Site The Gray Fossil Site is an Early Pliocene assemblage of fossils dating between 4.5 and 4.9 ...
'' 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 The red panda (''Ailurus fulgens''), also known as the lesser panda, is a small mammal native to the eastern Himalayas and southwestern China. It has dense reddish-brown fur with a black belly and legs, white-lined ears, a mostly white muzz ...
'' 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 The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
-age locality near Isère,
France France, officially the French Republic, is a country located primarily in Western Europe. Overseas France, Its overseas regions and territories include French Guiana in South America, Saint Pierre and Miquelon in the Atlantic Ocean#North Atlan ...
.


References


Further reading

* * * 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. * * Flynn, John J. Flynn, Michael A. Nedbal, J.W. Dragoo, and R.L. Honeycutt. (1998) "Whence the Red Panda?" ''Molecular Phylogenetics and Evolution''. Vol. 17, No. 2, November 2000, pp. 190–199

* Glatston, A.R. (1989). Talk Panda Biology. The Hague. * Glatston, A.R. (compiler) (1994). "The Red Panda, Olingos, Coatis, Raccoons, and their Relatives: Status survey and conservation action plan for Procyonids and Ailurids." * IUCN/SSC Mustelid, Viverrid, and Procyonid Specialist Group. IUCN/SSC, Gland, Switzerland. * * Hu, J.C. (1990). "Proceedings of studies of the red panda." Chinese Scientific Publishing, Beijing, China n Chinese * Wilson, Don E. and DeeAnn M. Reeder. (2005). Mammal of ''Species of the World''. Johns Hopkins University press. . {{Authority control Carnivorans Mammal families Extant Oligocene first appearances Taxa named by John Edward Gray