Procyonidae

{{Short description|Family of mammals}} {{EngvarB|date=June 2022}} {{Use dmy dates|date=June 2022}} {{Automatic taxobox | image = Procyonidae-01.jpg | image_caption = From top left to bottom right: raccoon (''Procyon''), ringtail (''Bassariscus''), South American coati (''Nasua''), northern olingo (''Bassaricyon''), kinkajou (''Potos'') | fossil_range = {{Fossil range|22.6|0}}Early Miocene to Holocene | taxon = Procyonidae | authority = Gray, 1825 | subdivision_ranks = Genera | type_genus = ''Procyon'' | type_genus_authority = Storr, 1780 | subdivision = *Extant genera: **''Bassariscus'' **''Procyon'' **''Nasua'' **''Nasuella'' **''Bassaricyon'' **''Potos'' *Extinct genera: **†''Arctonasua'' **†''Bassaricyonoides'' **†''Broiliana'' **†''Chapalmalania'' **†''Cyonasua'' **†''Edaphocyon'' **†''Parahyaenodon'' **†''Parapotos'' **†''Paranasua'' **†''Probassariscus'' **†''Protoprocyon'' **†''Stromeriella'' **†''Tetraprothomo'' | range_map = File:Procyonidae distribution.png | range_map_caption = Procyonidae distribution }}

'''Procyonidae''' ({{IPAc-en|ˌ|p|r|oʊ|s|iː|ˈ|ɒ|n|ɪ|d|iː}} {{respell|PROH|see|ON|i|dee}}) is a New World family of the order Carnivora.<ref name = MSW3>{{MSW3 Carnivora | id = 14001595 | pages = 624–628}}</ref> It includes the raccoons, ringtails, cacomistles, coatis, kinkajous, olingos, and olinguitos. Procyonids inhabit a wide range of environments and are generally omnivorous.

==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 family. This is apparent in their German 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 is a ''Honigbär'' (honey-bear). Dutch 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.<ref name="Flynn-2005">{{cite journal|last1=Flynn|first1=John|last2=Finarelli|first2=John|last3=Zehr|first3=Sarah|last4=Hsu|first4=Johnny|last5=Nedbal|first5=Michael|title=Molecular Phylogeny of the Carnivora (Mammalia): Assessing the Impact of Increased Sampling on Resolving Enigmatic Relationships|journal=Systematic Biology|volume=54|issue=2|year=2005|pages=317–337 |doi=10.1080/10635150590923326|pmid=16012099 |doi-access=free}}</ref> Procyonids share common morphological characteristics including a shortened rostrum, absent alisphenoid canals, and a relatively flat mandibular fossa.<ref>{{Cite journal |last1=Gompper |first1=Matthew E. |last2=Rega-Brodsky |first2=Christine C. |date=2016-01-06 |title=The Textbook in the Modern Mammalogy Classroom |journal=Journal of Mammalian Evolution |volume=23 |issue=2 |pages=217–218 |doi=10.1007/s10914-015-9319-4 |s2cid=17883651 }}</ref> Kinkajous have unique morphological characteristics consistent with their arboreally adapted locomotion, including a prehensile tail and unique femoral structure.<ref>{{cite thesis |title=The functional anatomy of prehensile and nonprehensile tails of the Platyrrhini (Primates) and Procyonidae (Carnivora) |first=Jason Michael |last=Organ |year=2007 |oclc=233832204 }}</ref><ref>{{Cite journal |last=Tarquini |first=Juliana |date=March 2021 |title=Femoral Shape in Procyonids (Carnivora, Procyonidae): Morphofunctional Implications, Size and Phylogenetic Signal |journal=Journal of Mammalian Evolution |volume=28 |issue=1 |pages=159–171 |doi=10.1007/s10914-019-09491-8 |s2cid=209332127 |hdl=11336/98644 |hdl-access=free }}</ref>

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 teeth, these are poorly developed in most species, especially the raccoons. Apart from the kinkajou, procyonids have the dental formula: {{DentalFormula|upper=3.1.4.2|lower=3.1.4.2}} for a total of 40 teeth. The kinkajou has one fewer premolar in each row: {{DentalFormula|upper=3.1.3.2|lower=3.1.3.2}} 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,<ref>{{Cite journal |last1=Gompper |first1=Matthew E. |last2=Gittleman |first2=John L. |last3=Wayne |first3=Robert K. |date=1997-04-01 |title=Genetic relatedness, coalitions and social behaviour of white-nosed coatis, Nasua narica |journal=Animal Behaviour |volume=53 |issue=4 |pages=781–797 |doi=10.1006/anbe.1996.0344 |s2cid=53170578 }}</ref> while kinkajous have been found to form social groups of two males and one female.<ref>{{Cite journal |last1=Kays |first1=Roland W. |last2=Gittleman |first2=John L. |date=April 2001 |title=The social organization of the kinkajou Potos flavus (Procyonidae) |journal=Journal of Zoology |volume=253 |issue=4 |pages=491–504 |doi=10.1017/S0952836901000450 }}</ref> 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.<ref>{{Cite web |last=Jirik |first=Kate |title=LibGuides: North American Ringtail (Bassariscus astutus) Fact Sheet: Reproduction & Development |url=https://ielc.libguides.com/sdzg/factsheets/ringtail/reproduction |access-date=2022-11-15 |website=ielc.libguides.com |language=en}}</ref><ref>{{Cite web |last=Jirik |first=Kate |title=LibGuides: Kinkajou (Potos flavus) Fact Sheet: Summary |url=https://ielc.libguides.com/sdzg/factsheets/kinkajou/summary-page |access-date=2022-11-15 |website=ielc.libguides.com |language=en}}</ref><ref>{{Cite web |title=Coati {{!}} San Diego Zoo Animals & Plants |url=https://animals.sandiegozoo.org/animals/coati |access-date=2022-11-15 |website=animals.sandiegozoo.org}}</ref><ref>{{Cite web |date=2012-02-07 |title=Raccoon Nation ~ Raccoon Facts {{!}} Nature |url=https://www.pbs.org/wnet/nature/raccoon-nation-raccoon-fact-sheet/7553/ |access-date=2022-11-15 |website=Nature |language=en-US}}</ref>

===Evolution===

Procyonid fossils once thought to belong to the genus ''Bassariscus'', which includes the modern ringtail and cacomistle, have been identified from the Miocene epoch, around 20 million years (Ma) ago. It has been suggested that early procyonids were an offshoot of the canids that adapted to a more omnivorous diet.<ref name="EoM">{{cite book |last=Russell |first=James |url=https://archive.org/details/encyclopediaofma00mals_0/page/98 |title=The Encyclopedia of Mammals |publisher=Facts on File |year=1984 |isbn=978-0-87196-871-5 |editor=Macdonald |editor-first=D. |location=New York |pages=[https://archive.org/details/encyclopediaofma00mals_0/page/98 98–99] |url-access=registration}}</ref> The recent evolution of procyonids has been centered on Central America (where their diversity is greatest);<ref name=ZooKeys/> they entered the formerly isolated South America as part of the Great American Interchange,<ref name="Koepfli2007">{{cite journal |last1=Koepfli |first1=K.-P. |last2=Gompper |first2=M. E. |last3=Eizirik |first3=E. |last4=Ho |first4=C.-C. |last5=Linden |first5=L. |last6=Maldonado |first6=J. E. |last7=Wayne |first7=R. K. |year=2007 |title=Phylogeny of the Procyonidae (Mammalia: Carvnivora): Molecules, morphology and the Great American Interchange |url=https://repository.si.edu/handle/10088/6026 |journal=Molecular Phylogenetics and Evolution |volume=43 |issue=3 |pages=1076–1095 |citeseerx=10.1.1.495.2618 |doi=10.1016/j.ympev.2006.10.003 |pmid=17174109|bibcode=2007MolPE..43.1076K |hdl=10088/6026 }}</ref> beginning about 7.3 Ma ago in the late Miocene, with the appearance of ''Cyonasua''.<ref name="Woodburne2010">{{cite journal|last=Woodburne|first=M. O.|author-link=Michael Woodburne|title=The Great American Biotic Interchange: Dispersals, Tectonics, Climate, Sea Level and Holding Pens|journal=Journal of Mammalian Evolution|volume= 17|issue=4|date= 2010-07-14|pages= 245–264|doi= 10.1007/s10914-010-9144-8|pmid=21125025|pmc=2987556}}</ref> Some fossil procyonids such as ''Stromeriella'' were also present in the Old World, before going extinct in the Pliocene.<ref>R. L. Carroll. 1988. Vertebrate Paleontology and Evolution</ref>

Genetic studies have shown that kinkajous are a sister group to all other extant procyonids; they split off about 22.6 Ma ago.<ref name="EizirikMurphy2010">{{cite journal |last1=Eizirik |first1=E. |last2=Murphy |first2=W. J. |last3=Koepfli |first3=K.-P. |last4=Johnson |first4=W. E. |last5=Dragoo |first5=J. W. |last6=Wayne |first6=R. K. |last7=O'Brien |first7=S. J. |date=2010-02-04 |title=Pattern and timing of diversification of the mammalian order Carnivora inferred from multiple nuclear gene sequences |journal=Molecular Phylogenetics and Evolution |volume=56 |issue=1 |pages=49–63 |doi=10.1016/j.ympev.2010.01.033 |pmc=7034395 |pmid=20138220|bibcode=2010MolPE..56...49E }}</ref> The clades leading to coatis and olingos on one branch, and to ringtails and raccoons on the other, separated about 17.7 Ma ago.<ref name=ZooKeys/> The divergence between olingos and coatis is estimated to have occurred about 10.2 Ma ago,<ref name=ZooKeys/> at about the same time that ringtails and raccoons parted ways.<ref name=ZooKeys/><ref name="Koepfli2007"/> The separation between coatis and mountain coatis is estimated to have occurred 7.7 Ma ago.<ref>{{cite thesis |id={{ProQuest|1999244770}} |last=Nunes Tsuchiya |first=Mirian Tieko |title=Evolutionary History of Procyonidae (Mammalia, Carnivora): Integrating Genomics, Morphology and Biogeographic Modeling |year=2017 }}</ref>

==Classification== {{see also|List of procyonids}} There has been considerable historical uncertainty over the correct classification of several members. The red panda was previously classified in this family, but it is now classified in its own family, the Ailuridae, based on molecular biology studies. The status of the various olingos was disputed: some regarded them all as subspecies of ''Bassaricyon gabbii'' before DNA sequence data demonstrated otherwise.<ref name=ZooKeys>{{cite journal |title=Taxonomic revision of the olingos (''Bassaricyon''), with description of a new species, the Olinguito |last1=Helgen |first1=K. M. |last2=Pinto |first2=M. |last3=Kays |first3=R. |last4=Helgen |first4=L. |last5=Tsuchiya |first5=M. |last6=Quinn |first6=A. |last7=Wilson |first7=D. |last8=Maldonado |first8=J.|journal= ZooKeys|issue= 324|date= 2013-08-15|pages= 1–83|doi= 10.3897/zookeys.324.5827 |pmid=24003317 |pmc=3760134|doi-access=free |bibcode=2013ZooK..324....1H }}</ref>

The traditional classification scheme shown below on the left predates the recent revolution in our understanding of procyonid phylogeny based on genetic sequence analysis. This outdated classification groups kinkajous and olingos together on the basis of similarities in morphology that are now known to be an example of parallel evolution; 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 {{as of|2013|lc=y}}.<ref name=ZooKeys/><ref name="Koepfli2007"/><ref name="EizirikMurphy2010"/> Genus ''Nasuella'' was not included in these studies, but in a separate study was found to nest within ''Nasua''.<ref name="split">{{cite journal |last1=Helgen |first1=K. M. |last2=Kays |first2=R. |last3=Helgen |first3=L. E. |last4=Tsuchiya-Jerep |first4=M. T. N. |last5=Pinto |first5=C. M. |last6=Koepfli |first6=K. P. |last7=Eizirik |first7=E. |last8=Maldonado |first8=J. E. |date=August 2009 |title=Taxonomic boundaries and geographic distributions revealed by an integrative systematic overview of the mountain coatis, ''Nasuella'' (Carnivora: Procyonidae) |url=https://repository.si.edu/handle/10088/8168 |journal=Small Carnivore Conservation |volume=41 |pages=65–74 |hdl=10088/8168 |access-date=2013-08-20}}</ref>

{{cladogram|clades={{Clade|style=width:35em; |label1=Procyonidae |1={{Clade |1={{Clade |1={{Clade |1=''Bassaricyon'' (olingos and olinguito) |2=''Nasua'' and ''Nasuella'' (coatis) }} |2={{Clade |1=''Procyon'' (raccoons) |2=''Bassariscus'' (ringtail and cacomistle) }} }} |2=''Potos'' (kinkajou) }}}}}}

* '''FAMILY PROCYONIDAE''' ** '''Subfamily Procyoninae''' (nine species in four genera) *** '''Tribe Procyonini''' **** '''Subtribe Procyonina''' ***** Raccoons, ''Procyon'' ****** Crab-eating raccoon, ''Procyon cancrivorus'' ****** Cozumel raccoon, ''Procyon pygmaeus'' ****** Common raccoon, ''Procyon lotor'' **** '''Subtribe Nasuina''' ***** ''Nasua'' ****** South American coati or ring-tailed coati, ''Nasua nasua'' ****** White-nosed coati, ''Nasua narica'' ***** ''Nasuella'' ****** Western mountain coati, ''Nasuella olivacea'' ****** Eastern mountain coati, ''Nasuella meridensis'' *** '''Tribe Bassariscini''' **** ''Bassariscus'' ***** Ringtail, ''Bassariscus astutus'' ***** Cacomistle, ''Bassariscus sumichrasti'' ** '''Subfamily Potosinae''' (five species in two genera) *** ''Potos'' **** Kinkajou, ''Potos flavus'' *** ''Bassaricyon'' **** Northern olingo or Gabbi's olingo, ''Bassaricyon gabbii'' **** Eastern lowland olingo, ''Bassaricyon alleni'' **** Western lowland olingo, ''Bassaricyon medius'' **** Olinguito, ''Bassaricyon neblina''

===Phylogeny===

Several recent molecular studies have resolved the phylogenetic relationships between the procyonids, as illustrated in the cladogram below.<ref name="Koepfli2007"/><ref name=ZooKeys/><ref name=split/><ref name="Law-2018">{{Cite journal|last1=Law|first1=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>

{{clade |style=font-size:90%;line-height:100% |label1=Procyonidae |1={{clade |label1=''Potos'' |1=''Potos flavus'' (kinkajou) |2={{clade |1={{clade |label1=''Procyon'' |sublabel1=(raccoons) |1={{clade |1=''Procyon cancrivorus'' (crab eating raccoon) |2=''Procyon lotor'' (common raccoon) 60 px |3=''Procyon pygmaeus'' (Cozumel raccoon) }} |label2=''Bassariscus'' |2={{clade |1=''Bassariscus sumichrasti'' (cacomistle) |2=''Bassariscus astutus'' (ringtail) }} }} |2={{clade |label1=''Bassaricyon'' |sublabel1=(olingos) |1={{clade |1={{clade |1={{clade |1=''Bassaricyon medius'' (western lowland olingo) 60 px |2=''Bassaricyon alleni'' (eastern lowland olingo) 60 px }} |2=''Bassaricyon gabbii'' (northern olingo) 60 px }} |2=''Bassaricyon neblina'' (olinguito) 60 px }} |label2=Nasuina |sublabel2=(coatis) |2={{clade |1={{clade |1=''Nasua nasua'' (ring-tailed coati) |2={{clade |1=''Nasua narica'' (white-nosed coati) |label2=''Nasuella'' |2={{clade |1=''Nasuella olivacea'' (western mountain coati) |2=''Nasuella meridensis'' (eastern mountain coati) }} }} }} }} }} }} }} }}

===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'' 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== {{reflist}}

==External links== *{{Wikispecies-inline}} *{{Commons category-inline}}

Category:Procyonidae Category:Mammal families Category:Extant Burdigalian first appearances Category:Taxa named by John Edward Gray

Procyonidae

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