Procyonidae

Procyonidae is a New World family of the order Carnivora.[2] It comprises the raccoons, coatis, kinkajous, olingos, olinguitos, ringtails, and cacomistles. Procyonids inhabit a wide range of environments and are generally omnivorous.

Procyonids
Temporal range: 22.6–0 Ma
Early Miocene to Holocene
Procyonidae
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Superfamily: Musteloidea
Family: Procyonidae
Gray, 1825
Type genus
Procyon
Genera

†Parahyaenodon[1]
†Tetraprothomo[1]
†Angustictis
Bassariscus
†Probassariscus
†Edaphocyon
†Arctonasua
†Cyonasua
†Amphinasua
†Chapalmalania
†Protoprocyon
†Paranasua
Procyon
Nasua
Nasuella
†Bassaricynoides
†Parapotos
Bassaricyon
Potos

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 names: 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 respectively. However, it is now believed that procyonids are more closely related to mustelids than to bears.[3]

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:

Dentition
3.1.4.2
3.1.4.2

for a total of 40 teeth. The kinkajou has one less premolar in each row:

Dentition
3.1.3.2
3.1.3.2

for a total of 36 teeth.

They are mostly solitary animals; mothers generally raise litters of up to four young on their own.[4]

Evolution

Procyonid fossils once believed 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.[4] The recent evolution of procyonids has been centered on Central America (where their diversity is greatest);[5] they invaded the formerly isolated South America as part of the Great American Interchange,[6] beginning about 7.3 Ma ago in the late Miocene, with the appearance there of Cyonasua.[7]

Genetic studies have shown that kinkajous are a sister group to all other extant procyonids; they split off about 22.6 Ma ago.[8] The clades leading to coatis and olingos on one hand, and to ringtails and raccoons on the other, separated about 17.7 Ma ago.[5] The divergence between olingos and coatis is estimated to have occurred about 10.2 Ma ago,[5] at about the same time that ringtails and raccoons parted ways.[5][6]

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.[5]

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 the recent molecular studies.[5][6][8] Genus Nasuella was not included in these studies, but in a separate study was found to nest within Nasua.[9]

Procyonidae  
 
 
 

Bassaricyon (olingos and olinguito)

 
 

Nasua and Nasuella (coatis)

 
 
 
 

Procyon (raccoons)

 
 

Bassariscus (ringtail and cacomistle)

 
 
 
 

Potos (kinkajou)

 
 

Phylogeny

Several recent molecular studies have resolved the phylogenetic relationships between the procyonids, as illustrated in the cladogram below.[6][5][9][10]

Procyonidae
Potos

Potos flavus (kinkajou)

 
 
 
Procyon
 

Procyon cancrivorus (crab eating raccoon)

 
 

Procyon lotor (common raccoon)

 
(raccoons)
Bassariscus
 

Bassariscus sumichrasti (cacomistle)

 
 

Bassariscus astutus (ringtail)

 
 
 
 
Bassaricyon
 
 
 

Bassaricyon medius (western lowland olingo)

 
 

Bassaricyon alleni (eastern lowland olingo)

 
 
 

Bassaricyon gabbii (northern olingo)

 
 
 

Bassaricyon neblina (olinguito)

 
(olingos)
Nasuina
 
 

Nasua nasua (ring-tailed coati)

 
 
 

Nasua narica (white-nosed coati)

 
Nasuella
 

Nasuella olivacea (western mountain coati)

 
 

Nasuella meridensis (eastern mountain coati)

 
 
 
 
(coatis)
 
 
 

References
  1. Forasiepi, Analía M.; Agustin G. Martinelli; Francisco J. Goin (2007). "Taxonomic revision of Parahyaenodon argentinus Ameghino and its implications for the knowledge of the Mio-Pliocene large carnivorous mammals of South America". Ameghiniana. 44 (1): 143–159.
  2. Wozencraft, W.C. (2005). "Order Carnivora". In Wilson, D.E.; Reeder, D.M (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. pp. 624–628. ISBN 978-0-8018-8221-0. OCLC 62265494.
  3. Flynn, John; Finarelli, John; Zehr, Sarah; Hsu, Johnny; Nedbal, Michael (2005). "Molecular Phylogeny of the Carnivora (Mammalia): Assessing the Impact of Increased Sampling on Resolving Enigmatic Relationships" (PDF). Systematic Biology. 54 (2): 317–337. doi:10.1080/10635150590923326. ISSN 1063-5157. PMID 16012099.
  4. Russell, James (1984). Macdonald, D. (ed.). The Encyclopedia of Mammals. New York: Facts on File. pp. 98–99. ISBN 978-0-87196-871-5.
  5. Helgen, K. M.; Pinto, M.; Kays, R.; Helgen, L.; Tsuchiya, M.; Quinn, A.; Wilson, D.; Maldonado, J. (2013-08-15). "Taxonomic revision of the olingos (Bassaricyon), with description of a new species, the Olinguito". ZooKeys (324): 1–83. doi:10.3897/zookeys.324.5827. PMC 3760134. PMID 24003317.
  6. K.-P. Koepfli; M. E. Gompper; E. Eizirik; C.-C. Ho; L. Linden; J. E. Maldonado; R. K. Wayne (2007). "Phylogeny of the Procyonidae (Mammalia: Carvnivora): Molecules, morphology and the Great American Interchange" (PDF). Molecular Phylogenetics and Evolution. 43 (3): 1076–1095. CiteSeerX 10.1.1.495.2618. doi:10.1016/j.ympev.2006.10.003. ISSN 1055-7903. PMID 17174109.
  7. Woodburne, M. O. (2010-07-14). "The Great American Biotic Interchange: Dispersals, Tectonics, Climate, Sea Level and Holding Pens". Journal of Mammalian Evolution. 17 (4): 245–264. doi:10.1007/s10914-010-9144-8. PMC 2987556. PMID 21125025.
  8. Eizirik, E.; Murphy, W. J.; Koepfli, K.-P.; Johnson, W. E.; Dragoo, J. W.; Wayne, R. K.; O’Brien, S. J. (2010-02-04). "Pattern and timing of diversification of the mammalian order Carnivora inferred from multiple nuclear gene sequences". Molecular Phylogenetics and Evolution. 56 (1): 49–63. doi:10.1016/j.ympev.2010.01.033. PMC 7034395. PMID 20138220.
  9. Helgen, K. M.; Kays, R.; Helgen, L. E.; Tsuchiya-Jerep, M. T. N.; Pinto, C. M.; Koepfli, K. P.; Eizirik, E.; Maldonado, J. E. (August 2009). "Taxonomic boundaries and geographic distributions revealed by an integrative systematic overview of the mountain coatis, Nasuella (Carnivora: Procyonidae)" (PDF). Small Carnivore Conservation. 41: 65–74. Retrieved 2013-08-20.
  10. 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.
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