Procyonid
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Procyonidae ( ) is a New World
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 ...
of the
order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...
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 ...
. It includes the
raccoons 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 ...
,
ringtail The ringtail (''Bassariscus astutus'') is a mammal of the raccoon family native to arid regions of North America. It is widely distributed and well-adapted to its distributed areas. It has been legally trapped for its fur. Globally, it is list ...
s,
cacomistle The cacomistle (; ''Bassariscus sumichrasti''), also spelled cacomixtle, is a primarily nocturnal, arboreal, omnivorous member of the carnivoran family Procyonidae (coatis, kinkajous and raccoons). Depending on the location, its preferred habi ...
s,
coati Coatis (from Tupí), also known as coatimundis (), are members of the family Procyonidae in the genera '' Nasua'' and '' Nasuella'' (comprising the subtribe Nasuina). They are diurnal mammals native to South America, Central America, Mexico, ...
s,
kinkajou The kinkajou ( /ˈkɪŋkədʒuː/ ''KING-kə-joo''; ''Potos flavus'') is a tropical rainforest mammal of the family Procyonidae related to olingos, coatis, raccoons, and the ringtail and cacomistle. It is the only member of the genus ''Pot ...
s,
olingo The genus ''Bassaricyon'' consists of small Neotropical procyonids, popularly known as olingos (), cousins of the raccoon. They are native to the rainforests of Central and South America from Nicaragua to Peru. They are arboreal and nocturnal, ...
s, and
olinguito The olinguito (''Bassaricyon neblina'') is a mammal of the raccoon family Procyonidae that lives in montane forests in the Andes of western Colombia and Ecuador. It was classified as belonging to a new species in 2013. The specific name ''nebli ...
s. Procyonids inhabit a wide range of environments and are generally
omnivorous An omnivore () is an animal that regularly consumes significant quantities of both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize ...
.


Characteristics

Procyonids are relatively small animals, with generally slender bodies and long tails, though the common raccoon tends to be bulky. Because of their general build, the Procyonidae are often popularly viewed as smaller cousins of 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. This is apparent in their
German German(s) may refer to: * Germany, the country of the Germans and German things **Germania (Roman era) * Germans, citizens of Germany, people of German ancestry, or native speakers of the German language ** For citizenship in Germany, see also Ge ...
name, ''Kleinbären'' (small bears), including the names of the species: a raccoon is called a ''Waschbär'' (washing bear, as it "washes" its food before eating), a coati is a ''Nasenbär'' (nose-bear), while a
kinkajou The kinkajou ( /ˈkɪŋkədʒuː/ ''KING-kə-joo''; ''Potos flavus'') is a tropical rainforest mammal of the family Procyonidae related to olingos, coatis, raccoons, and the ringtail and cacomistle. It is the only member of the genus ''Pot ...
is a ''Honigbär'' (honey-bear).
Dutch Dutch or Nederlands commonly refers to: * Something of, from, or related to the Netherlands ** Dutch people as an ethnic group () ** Dutch nationality law, history and regulations of Dutch citizenship () ** Dutch language () * In specific terms, i ...
follows suit, calling the animals ''wasbeer'', ''neusbeer'' and ''rolstaartbeer'' (curl-tail bear) respectively. However, it is now believed that procyonids are more closely related to mustelids than to bears. Procyonids share common morphological characteristics including a shortened rostrum, absent alisphenoid canals, and a relatively flat mandibular fossa. Kinkajous have unique morphological characteristics consistent with their arboreally adapted locomotion, including a prehensile tail and unique femoral structure. Due to their omnivorous diet, procyonids have lost some of the adaptations for flesh-eating found in their carnivorous relatives. While they do have
carnassial Carnassials are paired upper and lower teeth modified in such a way as to allow enlarged and often self-sharpening edges to pass by each other in a shearing manner. This adaptation is found in carnivorans, where the carnassials are the modified f ...
teeth, these are poorly developed in most species, especially the raccoons. Apart from the kinkajou, procyonids have the
dental formula Dentition pertains to the development of teeth and their arrangement in the mouth. In particular, it is the characteristic arrangement, kind, and number of teeth in a given species at a given age. That is, the number, type, and morpho-physiology ...
: for a total of 40 teeth. The
kinkajou The kinkajou ( /ˈkɪŋkədʒuː/ ''KING-kə-joo''; ''Potos flavus'') is a tropical rainforest mammal of the family Procyonidae related to olingos, coatis, raccoons, and the ringtail and cacomistle. It is the only member of the genus ''Pot ...
has one fewer
premolar The premolars, also called premolar Tooth (human), teeth, or bicuspids, are transitional teeth located between the Canine tooth, canine and Molar (tooth), molar teeth. In humans, there are two premolars per dental terminology#Quadrant, quadrant in ...
in each row: for a total of 36 teeth. Most members of Procyonidae are solitary; however, some species form groups. Coati females will form bands of 4 to 24 individuals that forage together, while kinkajous have been found to form social groups of two males and one female. Certain procyonids give birth to one offspring like ringtails, olingos, and kinkajous while raccoons and coatis give birth to litters that range in size from 2 to 6 offspring.


Evolution

Procyonid fossils once believed to belong to the genus ''
Bassariscus ''Bassariscus'' is a genus in the family Procyonidae. There are two extant species in the genus: the ringtail or ring-tailed cat (''B. astutus'') and the cacomistle (''B. sumichrasti''). Genetic studies have indicated that the closest relatives ...
'', which includes the modern ringtail and cacomistle, have been identified from the
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 ...
epoch, around 20 million years ( Ma) ago. It has been suggested that early procyonids were an offshoot of the
canids Canidae (; from Latin, ''canis'', "dog") is a biological family of caniform carnivorans, constituting a clade. A member of this family is also called a canid (). The family includes three subfamilies: the Caninae, and the extinct Borophaginae ...
that adapted to a more omnivorous diet. The recent evolution of procyonids has been centered on Central America (where their diversity is greatest); they entered the formerly isolated South America as part of the
Great American Interchange The Great American Biotic Interchange (commonly abbreviated as GABI), also known as the Great American Interchange and the Great American Faunal Interchange, was an important late Cenozoic paleozoogeographic biotic interchange event in which land ...
, beginning about 7.3  Ma ago in the late Miocene, with the appearance of ''
Cyonasua ''Cyonasua'' (meaning "dog-coati" in Greek) is an extinct genus of procyonid from the Late Miocene to Middle Pleistocene of South America. Fossils of ''Cyonasua'' have been found in Argentina ( Ituzaingó, Epecuén, Huayquerías, Monte Hermoso, ...
''. Some fossil procyonids such as '' Stromeriella'' were also present in the Old World, before going extinct in the Pliocene. Genetic studies have shown that kinkajous are a
sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
to all other extant procyonids; they split off about 22.6 Ma ago. The clades leading to coatis and olingos on one branch, and to ringtails and raccoons on the other, separated about 17.7 Ma ago. The divergence between olingos and coatis is estimated to have occurred about 10.2 Ma ago, at about the same time that ringtails and raccoons parted ways. The separation between coatis and mountain coatis is estimated to have occurred 7.7 Ma ago.


Classification

There has been considerable historical uncertainty over the correct classification of several members. 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 ...
was previously classified in this family, but it is now classified in its own family, the
Ailuridae 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 ...
, based on molecular biology studies. The status of the various
olingo The genus ''Bassaricyon'' consists of small Neotropical procyonids, popularly known as olingos (), cousins of the raccoon. They are native to the rainforests of Central and South America from Nicaragua to Peru. They are arboreal and nocturnal, ...
s was disputed: some regarded them all as
subspecies In Taxonomy (biology), biological classification, subspecies (: subspecies) is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (Morphology (biology), morpholog ...
of '' Bassaricyon gabbii'' before DNA sequence data demonstrated otherwise. The traditional classification scheme shown below on the left predates the recent revolution in our understanding of procyonid
phylogeny A phylogenetic tree or phylogeny is a graphical representation which shows the evolutionary history between a set of species or Taxon, taxa during a specific time.Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, M ...
based on genetic sequence analysis. This outdated classification groups kinkajous and olingos together on the basis of similarities in
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines *Morphology (archaeology), study of the shapes or forms of artifacts *Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...
that are now known to be an example of
parallel evolution Parallel evolution is the similar development of a trait in distinct species that are not closely related, but share a similar original trait in response to similar evolutionary pressure.Zhang, J. and Kumar, S. 1997Detection of convergent and pa ...
; similarly, coatis are shown as being most closely related to raccoons, when in fact they are closest to olingos. Below right is a cladogram showing the results of molecular studies . Genus ''Nasuella'' was not included in these studies, but in a separate study was found to nest within ''Nasua''. * FAMILY PROCYONIDAE ** Subfamily Procyoninae (nine species in four genera) *** Tribe Procyonini **** Subtribe Procyonina *****
Raccoons 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 ...
, ''Procyon'' ****** Crab-eating raccoon, ''Procyon cancrivorus'' ******
Cozumel raccoon The Cozumel raccoon (''Procyon pygmaeus'') is a critically endangered species of island raccoon endemic on Cozumel Island off the coast of the Yucatan Peninsula, Mexico. It is sometimes also called the pygmy raccoon, dwarf raccoon, Cozumel Isl ...
, ''Procyon pygmaeus'' ******
Common 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 ...
, ''Procyon lotor'' **** Subtribe Nasuina ***** ''
Nasua ''Nasua'' is a genus of coatis of the family Procyonidae. Two additional species of coatis, commonly known as mountain coatis, are placed in the genus '' Nasuella''. Characteristics ''Nasua'' differs from ''Nasuella'' in being larger and ...
'' ******
South American coati The South American coati (''Nasua nasua''), also known as the ring-tailed coati or brown-nosed coati, is a coati species and a member of the raccoon family ( Procyonidae), found in the tropical and subtropical parts of South America. An adult g ...
or ring-tailed coati, ''Nasua nasua'' ******
White-nosed coati The white-nosed coati (''Nasua narica''), also known as the coatimundi (), is a species of coati and a member of the family Procyonidae (raccoons and their relatives). Local Spanish names for the species include ''antoon'', ''gato solo'', ''pi ...
, ''Nasua narica'' ***** ''
Nasuella Mountain coatis are two species of procyonid mammals from the genus ''Nasuella''. Unlike the larger coatis from the genus '' Nasua'', mountain coatis only weigh and are endemic to the north Andean highlands in South America. Genetics and taxon ...
'' ******
Western mountain coati The western mountain coati or western dwarf coati (''Nasuella olivacea'') is a small procyonid, found in cloud forest and páramo at altitudes of in the Andes of Colombia and Ecuador.Helgen, K. M., R. Kays, L. E. Helgen, M. T. N. Tsuchiya-Jerep ...
, ''Nasuella olivacea'' ****** Eastern mountain coati, ''Nasuella meridensis'' *** Tribe Bassariscini **** ''
Bassariscus ''Bassariscus'' is a genus in the family Procyonidae. There are two extant species in the genus: the ringtail or ring-tailed cat (''B. astutus'') and the cacomistle (''B. sumichrasti''). Genetic studies have indicated that the closest relatives ...
'' *****
Ringtail The ringtail (''Bassariscus astutus'') is a mammal of the raccoon family native to arid regions of North America. It is widely distributed and well-adapted to its distributed areas. It has been legally trapped for its fur. Globally, it is list ...
, ''Bassariscus astutus'' *****
Cacomistle The cacomistle (; ''Bassariscus sumichrasti''), also spelled cacomixtle, is a primarily nocturnal, arboreal, omnivorous member of the carnivoran family Procyonidae (coatis, kinkajous and raccoons). Depending on the location, its preferred habi ...
, ''Bassariscus sumichrasti'' ** Subfamily Potosinae (five species in two genera) *** ''Potos'' ****
Kinkajou The kinkajou ( /ˈkɪŋkədʒuː/ ''KING-kə-joo''; ''Potos flavus'') is a tropical rainforest mammal of the family Procyonidae related to olingos, coatis, raccoons, and the ringtail and cacomistle. It is the only member of the genus ''Pot ...
, ''Potos flavus'' *** ''
Bassaricyon The genus ''Bassaricyon'' consists of small Neotropical procyonids, popularly known as olingos (), cousins of the raccoon. They are native to the rainforests of Central and South America from Nicaragua to Peru. They are arboreal and nocturnal, ...
'' ****
Northern olingo The northern olingo (''Bassaricyon gabbii''), also known as the bushy-tailed olingo or, simply, the olingo (due to it being the most common of the species), is an arboreal (tree-dwelling) member of the raccoon Family (taxonomy), family, Procyonid ...
or Gabbi's olingo, ''Bassaricyon gabbii'' **** Eastern lowland olingo, ''Bassaricyon alleni'' **** Western lowland olingo, ''Bassaricyon medius'' ****
Olinguito The olinguito (''Bassaricyon neblina'') is a mammal of the raccoon family Procyonidae that lives in montane forests in the Andes of western Colombia and Ecuador. It was classified as belonging to a new species in 2013. The specific name ''nebli ...
, ''Bassaricyon neblina''


Phylogeny

Several recent molecular studies have resolved the phylogenetic relationships between the procyonids, as illustrated in the cladogram below.


Extinct taxa

Below is a list of extinct taxa (many of which are fossil genera and species) compiled in alphabetical order under their respective subfamilies. * Procyonidae J.E. Gray, 1825 ** † Broilianinae Dehm, 1950 *** †'' Broiliana'' Dehm, 1950 **** †''B. dehmi'' Beaumont & Mein, 1973 **** †''B. nobilis'' Dehm, 1950 *** †'' Stromeriella'' Dehm, 1950 **** †''S. depressa'' Morlo, 1996 **** †''S. franconica'' Dehm, 1950 ** Potosinae Trouessart, 1904 *** †'' Parapotos'' J.A. Baskin, 2003 **** †''P. tedfordi'' J.A. Baskin, 2003 ** Procyoninae J.E. Gray, 1825 *** †'' Arctonasua'' J.A. Baskin, 1982 **** †''A. eurybates'' J.A. Baskin, 1982 **** †''A. fricki'' J.A. Baskin, 1982 **** †''A. floridana'' J.A. Baskin, 1982 **** †''A. gracilis'' J.A. Baskin, 1982 **** †''A. minima'' J.A. Baskin, 1982 *** †'' Bassaricyonoides'' J.A. Baskin & Morea, 2003 **** †''B. stewartae'' J.A. Baskin & Morea, 2003 **** †''B. phyllismillerae'' J.A. Baskin & Morea, 2003 *** ''Bassariscus'' Coues, 1887 **** †''B. antiquus'' Matthew & Cook, 1909 **** †''B. casei'' Hibbard, 1952 **** †''B. minimus'' J.A. Baskin, 2004 **** †''B. ogallalae'' Hibbard, 1933 **** †''B. parvus'' Hall, 1927 *** †'' Chapalmalania'' Ameghino, 1908 **** †''C. altaefrontis'' Kraglievich & Olazábal, 1959 **** †''C. ortognatha'' Ameghino, 1908 *** †''
Cyonasua ''Cyonasua'' (meaning "dog-coati" in Greek) is an extinct genus of procyonid from the Late Miocene to Middle Pleistocene of South America. Fossils of ''Cyonasua'' have been found in Argentina ( Ituzaingó, Epecuén, Huayquerías, Monte Hermoso, ...
'' Ameghino, 1885 ''Amphinasua'' Moreno & Mercerat, 1891; ''Brachynasua'' Ameghino & Kraglievich 1925; ''Pachynasua'' Ameghino, 1904**** †''C. argentina'' Ameghino 1885 **** †''C. argentinus'' (Burmeister, 1891) **** †''C. brevirostris'' (Moreno & Mercerat, 1891) ''Amphinasua brevirostris'' Moreno & Mercerat, 1891**** †''C. clausa'' (Ameghino, 1904) ''Pachynasua clausa'' Ameghino, 1904**** †''C. groeberi'' Kraglievich & Reig, 1954 ''Amphinasua groeberi'' Cabrera, 1936**** †''C. longirostris'' (Rovereto, 1914) **** †''C. lutaria'' (Cabrera, 1936) ''Amphinasua lutaria'' Cabrera, 1936**** †''C. meranii'' (Ameghino & Kraglievich 1925) ''Brachynasua meranii'' Ameghino & Kraglievich 1925**** †''C. pascuali'' Linares, 1981 ''Amphinasua pascuali'' Linares, 1981**** †''C. robusta'' (Rovereto, 1914) *** †'' Edaphocyon'' Wilson, 1960 **** †''E. lautus'' J.A. Baskin, 1982 **** †''E. palmeri'' J.A. Baskin & Morea, 2003 **** †''E. pointblankensis'' Wilson, 1960 *** ''Nasua'' Storr, 1780 **** †''N. pronarica'' Dalquest, 1978 **** †''N. mastodonta'' Emmert & Short, 2018 **** †''N. nicaeensis'' Holl, 1829 *** †'' Parahyaenodon'' Ameghino, 1904 **** †''P. argentinus'' Ameghino, 1904 *** †'' Paranasua'' J.A. Baskin, 1982 **** †''P. biradica'' J.A. Baskin, 1982 *** †'' Probassariscus'' Merriam, 1911 **** †''P. matthewi'' Merriam, 1911 *** ''Procyon'' Storr, 1780 **** †''P. gipsoni'' Emmert & Short, 2018 **** †''P. megalokolos'' Emmert & Short, 2018 **** †''P. rexroadensis'' Hibbard, 1941 *** †'' Protoprocyon'' Linares, 1981 ''Lichnocyon'' J.A. Baskin, 1982**** †''P. savagei'' Linares, 1981 ''Lichnocyon savagei'' J.A. Baskin, 1982*** †'' Tetraprothomo'' Ameghino, 1908 **** †''T. argentinus'' Ameghino, 1908


References


External links

* * {{Authority control Mammal families Extant Burdigalian first appearances Taxa named by John Edward Gray