Taxonomy and Systematics
The Procyonidae (raccoon family) includes around a dozen species in six genera. Because the members of this family are diverse in appearance, they were not immediately recognized by early naturalists as a coherent group, causing some confusion. For example, the Northern Raccoon (Procyon lotor) was described first as a fox and then a bear and the Kinkajou (Potos flavus) was originally described as a primate (a lemur), based in part on its forward-facing eyes (Kays 2009).
Kays (2009) recognized the following six genera and 12 species of procyonids:
Bassaricyon (3 species [but now 4]): Lowland Olingo (B. alleni), Northern Olingo (B. gabbii), and a then undescribed species, from the Andean cloud forests of Colombia and Ecuador, that has since been described (Helgen et al. 2013) as the Olinguito (B. neblina). Helgen et al. (2013) also recognized a fourth species, B. medius. They concluded that B. gabbii occurs in Central America and that the Lowland Olingo actually includes two species, B. alleni on the eastern side of the Andes and B. medius on the western side.
Bassariscus (2 species): Ringtail (B. astutus), found on the west coast of the United States through the southwest and into northern Mexico, and Cacomistle (B. sumichrasti), found in Central America and southern Mexico.
Nasua (2 species): South American Coati (Nasua nasua), found over much of South America except for the southern third, and White-nosed Coati (Nasua narica), found throughout Central America and into the southwestern United States. The Dwarf (or Cozumel) Coati, from Cozumel Island, Mexico, is sometimes recognized as a distinct species, N. nelsoni.
Nasuella (1 species [but now 2]): Mountain Coati (Nasuella olivacea), found in montane forests above 2000 meters in northwestern South America. Helgen et al. (2009) suggested that this might be the least studied carnivore genus globally. Morphological and molecular phylogenetic investigations by Helgen et al. led them to the conclusion that there are actually two distinct mountain coati species, which they referred to as the Eastern Mountain Coati (Nasuella meridensis), found in the Venezuelan Andes and formerly recognized as a subspecies, and the Western Mountain Coati (N. olivacea), found in the Andes of Colombia and Ecuador. Their molecular phylogenetic analyses also suggested that Nasuella may fall within the genus Nasua as the sister lineage to Nasua narica, making the genus Nasua paraphyletic as currently defined, although a strong conclusion on this question will require additional morphological and genetic comparisons, including data from independent genetic markers. If this conclusion stands, however, the genus name Nasuella would likely be relegated to a synonym of Nasua and all the coatis and mountain coatis would be included in the genus Nasua..
Potos (1 species): Kinkajou (Potos flavus), found from southern Mexico through the northern half of South America east of the Andes.
Procyon (3 species): Crab-eating Raccoon (P. cancrivorus), found from southern Central America through the northern two thirds of South America east of the Andes; Northern Raccoon (Procyon lotor), found from southern Canada through Central America (and established as an invasive species in Europe, Russia, and Japan); and Cozumel Raccoon (P. pygmaeus), from Cozumel Island, Mexico. Dozens of supposedly distinct forms of Northern Raccoon have been described over the years, but based on more recent work these are not considered to be valid. Interestingly, one result is that raccoons on a number of Caribbean islands that were formerly viewed as several distinct endemic species worthy of special conservation efforts are now viewed as invasive pests from the mainland that threaten local biodiversity. Similarly, the extinction of the Barbados Raccoon (last recorded in 1964) is now viewed as a stroke of good luck for the biodiversity of Barbados rather than a sad loss. The raccoons on Cozumel Island are believed to have arrived there long before humans and are still generally recognized as a distinct species, athough this could change once variation in mainland forms is studied more thoroughly.
Procyonids are medium-sized carnivorans, ranging from around 1 to 10 kg. Only the coatis show significant sexual dimorphism, with males being larger. All procyonids can climb well, but some are far better adapted than others for arboreal life: raccoons and coatis frequently climb trees for food or safety but rarely move among tree canopies, whereas the Kinkajou is one of the most arboreally specialized species in the order Carnivora, regularly moving from treetop to treetop in the dark, covering 2 km in a night. Kinkajous have fully reversible hindfeet, allowing them to descend branches and hang, and have remarkable vertebral flexibility. The Kinkajou and the Binturong (family Viverridae) are the only carnivorans with a prehensile tail, from which they often hang to reach fruit. Kinkajous are also unusual in having a fully convergent grasp, i.e., they are able to hold objects (such as food) with one hand.
Most procyonids are habitat generalists, although the Olinguito and mountain coatis are Andean cloud forest specialists. Procyonids are generally associated with trees and any forest across most of the Americas is likely to be home to at least one procyonid (as many as six species may be found in some parts of Panama). Kinkajous occur in virtually all tropical forest types from lowland tropical rainforest to high elevation cloud forest to dry forest. White-nosed and South American Coatis also use a wide variety of forest types, but move into drier chaco, cerrado, and mountainous deserts as well. Raccoons are tied to the aquatic environments they feed in. They find abundant habitat in wet forests, but also follow waterways and shorelines into drier habitats. In addition, Northern Raccoons thrive in suburban and even urban habitats (especially since the 1970s and 1980s). Ringtails are found in a range of forest habitat types, but are less tied to forest than other procyonids, often occurring in rocky deserts with cliffs. Cacomistles are found only in rainy tropical areas, but use a range of habitats, including mature oak cloud forest, secondary forest, and overgrown pastures. Some procyonids actually seem to benefit from human activity (e.g., Northern Raccoons and, to a lesser extent, Ringtails) and Cacomistles, South American Coatis, White-nosed Coatis, and Kinkajous seem to tolerate moderate forest fragmentation reasonably well, so long as they are not heavily hunted. Some other procyonids, such as mountain coatis and Olinguitos are believed to be far more sensitive to habitat disturbance, although little information is available for these taxa.
Procyonids are among the least carnivorous of all families in the Carnivora. Northern Raccoons are extreme omnivores and Kinkajous are surely among the most frugivorous mammals. Data from Panama, Venezuela, and French Guiana indicated that the Kinkajou diet is up to 99% fruit, although a study in Bolivia found substantial amounts of ants and ant nest material in the stomachs of six Kinkajous.
The most arboreal procyonids (Kinkajous, olingos, Cacomistles) make loud calls that probably function to communicate over long distances. This is unusual in the Carnivora, although common for arboreal primates.
Most procyonids are generally nocturnal, spending the day sleeping in a tree cavity or rock den and only emerging after sunset. Coatis, however, sleep in trees during the night and are active during the day. Coatis are also the most social procyonids. Females and their young travel in groups of up to 65 individuals, although smaller groups of 10 to 30 are more common. Adult male coatis (especially South American Coatis) sometimes trail along, but are more likely to forage alone. Kinkajous and Northern Raccoons have also been found to exhibit some degree of sociality, at least under some circumstances.
Because most procyonids are nocturnal, and often arboreal, they tend not to intreract much with humans. Although in the tropics they may take some fruit from gardens, they tend to be viewed more as charming visitors than pests (Northern Raccoons raiding suburban and urban garbage cans are more often viewed as pests). Starting in the 1980s, a new strain of raccoon rabies spread rapidly through high density populations in the eastern United States, reducing urban raccoon populations but also threatening other wildlife and even humans. At times, there has been a significant trade in raccoon pelts, ranging from 400,000 to two million over most of the 20th century, but peaking at a harvest of 5.1 million in the winter of 1979/80.
Although most procyonid species are not threatened at this point in time, the Cozumel Raccoon and Cozumel (or Dwarf ) Coati are highly endangered (Kays  treated the former as a valid raccoon species and the latter as a subspecies of the White-nosed Coati, but he noted that the Cozumel Coati is sometimes treated as a full species). The Olinguito has a small range in a region with intense conversion of forest to agriculture; the mountain coatis also have small ranges and may be threatened by conversion of their forest habitat to agriculture.
(Kays 2009 and references therein)
- Fulton, T.L. and C. Strobeck. 2007. Novel phylogeny of the raccoon family (Procyonidae: Carnivora) based on nuclear and mitochondrial DNA evidence. Molecular Phylogenetics and Evolution 43: 1171-1177.
- Helgen, K.M., R. Kays, L.E. Helgen, et al. 2009. Taxonomic boundaries and geographic distributions revealed by an integrative systematic overview of the mountain coatis, Nasuella (Carnivora: Procyonidae). Small Carnivore Conservation 41: 65-74.
- Kays, R. 2009. Family Procyonidae (Raccoons). Pp. 504-530 in: Wilson, D.E. & Mittermeier, R.A., eds. Handbook of the Mammals of the World. Volume 1. Carnivores. Lynx Edicions, Barcelona.
- Koepfli, K.P., M.E. Gompper, E. Eizirik, et al. 2007. Phylogeny of the Procyonidae (Mammalia: Carnivora): molecules, morphology, and the Great American Interchange. Molecular Phylogenetics and Evolution 43: 1076–1095.
Molecular Biology and Genetics
Statistics of barcoding coverage
|Specimen Records:||264||Public Records:||44|
|Specimens with Sequences:||210||Public Species:||9|
|Specimens with Barcodes:||102||Public BINs:||7|
|Species With Barcodes:||8|
Procyonidae is a New World family of the order Carnivora. It includes the raccoons, coatis, kinkajous, olingos, olinguitos, ringtails and cacomistles. Procyonids inhabit a wide range of environments and are generally omnivorous.
Procyonids are relatively small animals, with generally slender bodies and long tails (though the common raccoon tends to be bulky). Many procyonids have banded tails, and distinctive facial markings – these are especially visible in the raccoons. Like bears, procyonids are plantigrade, walking on the soles of their feet. Most species have non-retractile claws.
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 he "washes" his 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.
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:
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. The recent evolution of procyonids has been centered in Central America (where their diversity is greatest); they invaded formerly isolated South America as part of the Great American Interchange, beginning about 7.3 Ma ago in the late Miocene, with the appearance there of Cyonasua.
Genetic studies have shown that kinkajous are a sister group to all other extant procyonids; they split off about 22.6 Ma ago. The clades leading to coatis and olingos on one hand, 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.
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 it 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.
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 to the right is a cladogram showing the results of the recent 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
- Subtribe Nasuina
- Tribe Bassariscini
- Tribe Procyonini
- Subfamily Potosinae (five species in two genera)
- Subfamily Procyoninae (nine species in four genera)
- Wozencraft, W. C. (2005). "Order Carnivora". In Wilson, D. E.; Reeder, D. M. Mammal Species of the World (3rd ed.). Johns Hopkins University Press. ISBN 978-0-8018-8221-0. OCLC 62265494.
- Russell, James (1984). Macdonald, D., ed. The Encyclopedia of Mammals. New York: Facts on File. pp. 98–99. ISBN 0-87196-871-1.
- 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.
- 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". Molecular Phylogenetics and Evolution 43 (3): 1076–1095. doi:10.1016/j.ympev.2006.10.003. PMID 17174109.
- 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.
- 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.
- 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)". Small Carnivore Conservation 41: 65–74. Retrieved 2013-08-20.