White-headed Capuchin

Taxobox
name = White-headed CapuchinMSW3 Groves|pages=137|id=12100270]
status = LC
status_system = iucn3.1
status_ref = IUCN2008|assessors=Causado, J., Cuarón, A.D., Shedden, A., Rodríguez-Luna, E. & de Grammont, P.C.|year=2008|id=40020|title=Cebus capucinus|downloaded=7 October 2008]

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image_width = 200px
regnum = Animalia
phylum = Chordata
classis = Mammalia
ordo = Primates
familia = Cebidae
genus = "Cebus"
species = "C. capucinus"
binomial = "Cebus capucinus"
binomial_authority = (Linnaeus, 1758)
synonyms =
* "albulus" (Pusch, 1942)
* "curtus" (Bangs, 1905)
* "hypoleucus" (É. Geoffroy, 1812)
* "imitator" (Thomas, 1903)
* "limitaneus" (Hollister, 1914)
* "nigripectus" (Elliot, 1909)

The White-headed Capuchin ("Cebus capucinus"), also known as the White-faced Capuchin or White-throated Capuchin, is a medium-sized New World monkey of the family Cebidae, subfamily Cebinae. Native to the forests of Central America and the extreme north-western portion of South America, White-throated Capuchins are important to rainforest ecology by their role in dispersing seeds and pollen.

Among the best known monkeys, the White-headed Capuchin is recognized as the typical companion to the organ grinder. In recent years the species has become popular in North American media. It is a highly intelligent monkey and has been trained to assist for paraplegic persons.

It is a medium-sized monkey, weighing up to convert|3.9|kg|lb|lk=on|abbr=on. It is mostly black, but with a pink face and white on much of the front part of the body, giving it its common name. It has a distinctive prehensile tail that is often carried coiled up and is used to help support the monkey when it is feeding beneath a branch.

In the wild, White-headed Capuchins are versatile, living in many different types of forest, and eating many different types of food, including fruit, other plant material, invertebrates, and small vertebrates. They live in groups that can exceed 20 animals and include both males and females. They are noted for their tool use, including rubbing plants over their fur in an apparent use of herbal medicine, and also using tools as weapons and for getting to food. They are long-lived monkeys, with a maximum recorded age of over 54 years.

Description

Like other monkeys in the genus "Cebus", the White-headed Capuchin is named after the order of Capuchin friars: the cowls worn by these friars closely resemble the monkey's head coloration. [cite web|title=Capuchin Franciscans|url=http://capuchinfranciscans.org/sub_faq.html|accessdate=2008-09-01] cite book|title=The Natural History of Costa Rican Mammals|author=Wainwright, M.|year=2002|page=135–139|isbn=0-9705678-1-2] The White-headed Capuchin has mostly black fur, with white to yellowish fur on the neck, throat, chest, shoulders, and upper arms.cite book|title=Neotropical Rainforest Mammals A Field Guide|edition=Second Edition|author=Emmons, L.|page=130–131|year=1997|isbn=0-226-20721-8] The face is pink. A V-shaped area of black fur on the crown of the head is distinctive.cite book|title=The Pictorial Guide to the Living Primates|aithor=Rowe, N.|page=95|year=1996|isbn=0-9648825-0-7] It has a prehensile tail that is often held coiled, giving the White-headed Capuchin the nickname "ringtail". [cite web|title=Medical Dictionary Capuchin Monkey|url=http://medical-dictionary.thefreedictionary.com/Capuchin+(monkey)|accessdate=2008-09-01]

Adults reach a length of convert|335|and|453|mm|in|lk=on|abbr=on, excluding tail, and a weight of up to convert|3.9|kg|lb|lk=on|abbr=on. The tail is prehensile, dextrous, and longer than the body, at up to convert|551|mm|in|abbr=on in length. Males are about 27% larger than females.cite book|title=Primates in Perspective|chapter=The Cebines|author=Jack, K.|editor = Campbell, C., Fuentes, A., MacKinnon, K., Panger, M., & Bearder, S.|year=2007|page=107–120|isbn=978-0-19-517133-4] The brain of a White-headed Capuchin is about convert|79.2|g|oz|lk=on|abbr=on.

Distribution and habitat

[
Manuel Antonio National Park, Costa Rica]

The White-headed Capuchin is found in much of Central America and a small portion of South America. In Central America, its range includes much of Honduras, Nicaragua, Costa Rica and Panama.cite book|title=New Perspectives in the Study of Mesoamerican Primates|chapter=Taxonomy and Distributions of Mesoamerican Primates|author=Rylands, A., Groves, C., Mittermeier, R., Cortes-Ortiz, L., and Hines, J.|year=2006|page=40–43|isbn=0-387-25854-X] It has also been reported to occur in eastern Guatemala and southern Belize, but these reports are unconfirmed. In South America the White-headed Capuchin is found in the extreme north-western strip between the Pacific Ocean and the Andes Mountains in Colombia and northwestern Ecuador.

They are found in many different types of forest, including mature and secondary forests, and including evergreen and deciduous forests, dry and moist forests, and mangrove and montane forests.cite book|title=Field Guide to the Wildlife of Costa Rica|author=Henderson, C.|year=2000|page=454–455|isbn=0-292-73459-X] But they appear to prefer primary or advanced secondary forests. And higher densities of White-headed Capuchins appear to be found in older areas of forest and in areas containing evergreen forest, as well as areas with more water availability during the dry season. [cite book | author = DeGama, H. and Fedigan, L. | editor = Estrada, A.; Garber, P.A.; Pavelka, M.S.M.; Luecke, L. | chapter = The Effects of Forest Fragment Age, Isolation, Size, Habitat Type, and Water Availability on Monkey Density in a Tropical Dry Forest | title = New Perspectives in the Study of Mesoamerican Primates| page = 165–186 | isbn=978-0-387-25854-6 | year = 2006]

The White-headed Capuchin is among the most commonly seen monkeys in Central America's national parks, such as Manuel Antonio National Park, Corcovado National Park, Santa Rosa National Park and Soberania National Park. [cite book|title=Watching Wildlife Central America|author=Hunter, L. & Andrew, D.|year=2002|page=97, 100, 110, 130|isbn=1-86450-034-4]

Behavior

ocial structure

The White-headed Capuchin is a diurnal and arboreal animal. It moves primarily by walking on all four limbs. [cite book|title=New Perspectives in the Study of Mesoamerican Primates|chapter=Ontogenetic Influences on Positional Behavior in "Cebus" and "Alouatta"|author=Bezanson, L.|year=2006|page=333–344|isbn=0-387-25854-X] It forms troops of up to 24 monkeys with more females than males. On average there are 0.71 males for every female. The average troop size is 16 monkeys.

The troop is led by a dominant male and female. The females are generally related, as females tyically remain in their natal troop for life. Although the males are not necessarily related to each other, they are less competitive with each other than South American capuchin monkeys and they associate with each other and cooperate with each other to drive away predators. This may because, unlike in South American capuchins, the dominant male in the troop does not necessarily dominate breeding. Troop males tend to emigrate to new troops together. Male emigration to a new troop typically occurs about every 4 years.cite book|title=New Perspectives in the Study of Mesoamerican Primates|chapter=Dominance and Reproductive Success in Wild White-Faced Capuchins|author=Jack, K. and Fedigan, L.|year=2006|page=367–382|isbn=0-387-25854-X]

White-headed Capucin troops occupy home ranges of between convert|32|and|86|ha|acre|abbr=off|lk=on square metres. They cover between convert|1|and|3|km|mi|abbr=off|lk=on daily, averaging convert|2|km|mi|abbr=off|lk=on per day.cite book|title=Primates of Colombia|author=Defler, T.|page=237–235|year=2004|isbn=1-881-17383-6] Although they engage in activity that has been described as "territorial", more recent research indicates that White-headed Capuchin groups tend to behave aggressively to other conspecific groups regardless of where they meet, and that the aggression is not necessarily intended to exclude the conspecifics from a specific home range. [cite book|title=The Complete Capuchin|chapter=Behavioral Ecology|author=Fragaszy, D., Visalberghi, E., & Fedigan, L.|year=2004|page=38–39|isbn=0521667682]

Grooming pervades social interaction. Lower ranking animals are more likely to groom higher ranking animals than vice versa.cite book|title=The Complete Capuchin|chapter=Social Interactions, Relationships and Social Structure|author=Fragaszy, D., Visalberghi, E., & Fedigan, L.|year=2004|page=202–220|isbn=0521667682] Females do more grooming than males, and groom other females more than they groom males. Females associate closely among themselves. When non-group males attempt to supplant the existing group males, the females generally keep themselves and their infants away from any resulting fighting. However, the females sometimes form "coalitions" to attempt to help repel the intruders.

White-headed Capuchins sometimes interact with other sympatric monkey species. White-headed Capuchins sometimes travel with and even groom Geoffroy's Spider Monkeys. However, aggressive interactions between the capuchins and spider monkeys also occur.cite journal|title=Interspecific Interactions between "Cebus capucinus" and other Species: Data from Three Costa Rican Sites|author=Rose, L., Perry, S., Panger, M., Jack, K., Manson, J., Gros-Louis, J., and Mackinnin, K.|journal=International Journal of Primatology|volume=24|issue=4|date=August 2003|url=http://people.ucsc.edu/~evogel/cv/Roseetal2003.pdf|pages=780–785] Interactions between the White-headed Capuchin and Mantled Howler are infrequent, and sometimes result in the capuchins threatening the larger howlers. However, affiliative associations between the capuchins and howlers do sometimes occur, mostly involving juveniles playing together.

Although South American species of capuchins often travel with and feed together with squirrel monkeys, the White-headed Capuchin only rarely associates with the Central American Squirrel Monkey.cite book|title=On the Move|chapter=Social Manipulation Within and Between Troops Mediates Primate Group Movement|author=Boinski, S.|editor=Boinski, S. and Garber, P.|page=447–448|year=2000|isbn=0-226-06340-2] This may be because the distribution of food in the areas where Central American Squirrel Monkeys occur and is such that the cost to the squirrel monkeys of associating with the capuchins in lost feeding opportunities would not offset the benefit to the squirrel monkeys in predator detection. The minimal dietary overlap between the White-headed Capuchin and Central American Squirrel Monkey may also contribute to the minimal interaction. [cite book|title=The Complete Capuchin|chapter=Community Ecology|author=Fragaszy, D., Visalberghi, E., & Fedigan, L.|year=2004|page=70|isbn=0521667682]

Several non-primate animal species tend to follow groups of White-faced Monkeys or are otherwise attracted by their presence. White-lipped Peccaries and Common Agoutis are attracted by feeding White-headed Capuchins, looking for fruit that the capuchins drop. Several species of bird are also known to follow White-headed Capuchins looking for food. These include the Double-toothed Kite, the White Hawk and the Sharp-shinned Hawk.

Diet

White-headed Capuchins are omnivores with varied diets. Their primary foods are fruit and insects. They forage at all levels of the forest, and also forage on the ground. The prehensile tail assists with feeding, helping support the monkey when foraging for food below the branches.

Fruit can make up between 1/2 and 2/3 of the capuchins' diet. In one study in Panama they were noted eating 95 different fruit species. Among their favorite fruits are figs from the family Moraceae, mangos and related fruits from the family Anacardiaceae, the bean-like fruits from the family Leguminosae and fruits from the family Rubiaceae.cite book|title=The Complete Capuchin|chapter=Behavioral Ecology|author=Fragaszy, D., Visalberghi, E., & Fedigan, L.|year=2004|page=43–47|isbn=0521667682] They generally only eat the ripe fruit, testing for ripeness by smelling, tasting and prodding the fruit. They typically eat only the pulp and juice, spitting out the seeds and fibers. Other plant matter eaten includes flowers, young leaves, seeds of certain plants, and bromeliads. [cite book|title=New Perspectives in the Study of Mesoamerican Primates|chapter=Food Choice by Juevenile Capuchin Monkeys|author=MacKinnon, K.|year=2006|page=354–360|isbn=0-387-25854-X] They also drink water that gets trapped in the bromeliads.

Insect prey eaten includes beetle larvae, butterfly and moth caterpillars, ants, wasps, and ant and wasp larvae. They also eat larger prey. Such prey includes birds, bird eggs, frogs, lizards, crabs, mollusks and small mammals.cite video|title=Life of Mammals|people=David Attenborough|year2=2003|publisher=BBC Video] The population in Guanacaste, Costa Rica in particular is noted for hunting squirrels, magpies, White-crowned Parrots and baby coatis. The amount of vertebrate prey eaten varies by troop.

The diet can vary between the rainy and dry season. For example, in Guanacaste, Costa Rica the White-headed Capuchins can eat a wide variety of fruits and also caterpillars in the early rainy season (June to November). But in the dry season, only figs and a few other types of fruit are available. During the dry season, chitinous insects, ant and wasp larvae and vertebrates become a particularly important part of the White-headed Capuchin's diet.

During the dry season, access to water can also become an issue. White-headed Capuchins like to drink daily. In forests where water holes dry up during the dry season, there can be competition between groups over access to the remaining water holes.

Tool use

Capuchins are considered among the most intelligent of the New World monkeys; they have been the subject of many behavioural and intelligence studies. The capuchins' intelligence is thought to be a result of their feeding habits; they rely on ephemeral food sources which may be hard to find. In one particular study conducted in 2007, capuchins were found to be among the ten most intelligent primates, second to spider monkeys among New World monkeys.cite web|title=Chimps Knocked Off Top of the IQ Tree|url=http://www.timesonline.co.uk/tol/news/uk/article1654998.ece|author=Leake, D. & Dobson, R.|date=April 15, 2007|year=2007|accessdate=2008-09-01] The White-headed Capuchin is known to rub parts of certain plants into their fur. Plants used in this manner include citrus fruits, vines of the genera "Piper" and "Clematis", monkey comb (genus "Sloanea"), dumb cane and custard apple. Ants and millipedes are also used in this way. It is not definitively known what this fur rubbing is for. But this may deter parasites such as ticks and insects, or it may serve as a fungicide or bactericide or anti-inflammatory agent. Alternatively, it may be a form of scent marking.

White-headed Capuchins also use tools in other ways. They have been known to beat snakes with sticks in order to protect themselves or to get them to release an infant. In captivity, they have been known to use tools to get to food or to defend themselves, and in one case a White-headed Capuchin used a squirrel monkey as a projectile, hurling it at a human observer. Some populations also use trees or other hard surfaces as anvils in order to crack mollusks. And they sometimes use sticks as probes to explore openings. [cite book|title=The Complete Capuchin|chapter=Capuchins Use Objects as Tools|author=Fragaszy, D., Visalberghi, E., & Fedigan, L.|year=2004|page=173–183|isbn=0521667682]

The White-headed Capuchin's intelligence and ability to use tools allows them to be trained to assist paraplegics.cite web|title=Bronx Zoo: Monkey House - White-throated Capuchin|url=http://www.flickr.com/photos/wallyg/469410855/|accessdate=2008-09-01] Other species of capuchin monkey are also trained in this manner. [Citation|title=Monkeys as Helpers To Quadriplegics At Home |author=Blumenthal, D.|year=1987|date=1987-06-17|newspaper=The New York Times] White-headed Capuchins can also be trained for roles on television and movies, such as Marcel on the television series "Friends" They were also traditionally used as organ grinder monkeys.

Communication

The White-headed Capuchin is characterised as being noisy. Loud calls, such as barks and coughs, are used to communicate threat warnings, and softer calls, such as squeals, are used in intimate discourse. Facial expressions and scent are also important to communication. They sometimes engage in a practice known as "urine washing", in which a monkey rubs urine on its feet. [cite book|title=The Complete Capuchin|chapter=The Body|author=Fragaszy, D., Visalberghi, E., & Fedigan, L.|year=2004|page=102|isbn=0521667682] The exact purpose of this practice is unknown, but it may be a form of olfactory signal.

Reproduction

White-headed Capuchins use a polygamous mating system in which a male may mate with multiple females. Although the dominant male does not monopolize breeding, studies have shown that the dominant male does tend to father most of the young. This may be because the dominant male is more likely to copulate when the female is at peak fertility. [cite book|title=New Perspectives in the Study of Mesoamerican Primates|chapter=Post-conceptive Mating in White-Faced Capuchins|author=Carnegie, S., Fedigan, L., and Ziegler, T.|year=2006|page=387–405|isbn=0-387-25854-X]

Copulation takes about 2 minutes. Gestation period is 5 to 6 months. Usually a single young is born, but twins occur occasionally. Most births occur during the dry season from December to April.

The infant is carried across its mother's back for about 6 weeks. After about 7 to 8 weeks it can stray from its mother for brief periods. By about 6 months it can move around independently. Weaning occurs between 6 and 12 months. Females give birth every 26 months, on average. While the mother rests, the young spends most of its time playing, either on its own or with other juveniles.

Like other capuchin species, White-headed Capuchins mature slowly. Sexual maturity can be reached at 3 years. But on average, females give birth for the first time at 7 years old. Males attain reproductive maturity at 10 years old. White-headed Capuchins can also have long life spans given their size. Maximum recorded life span in captivity is over 54 years.

Conservation status

The White-headed Capuchin is regarded as "least concern" from a conservation standpoint by IUCN. However, their numbers are impacted by the fact that they are sometimes captured as pets. Their status can also be harmed by deforestation. However, deforestation may impact their main predator, the Harpy Eagle, more than it directly impacts the White-headed Capuchin, and so on a net basis deforestation may not be as harmful to the capuchin's status. White-headed Capuchin can adapt to forest fragmentation better than other species due to their ability to live in a wide variety of forest types and exploit a wide variety of food sources. [cite book|title=New Perspectives in the Study of Mesoamerican Primates|chapter=Concluding Comments and Conservation Priorities|author=Garber, P., Estrada, A., and Pavelka, M.|year=2006|page=570–571|isbn=0-387-25854-X] White-headed Capuchins are important to their ecosystems for a number of reasons, but especially on their capacity as seed and pollen dispersers.

Taxonomy

The White-headed Capuchin was one of the many species originally described by Linnaeus in his 18th century work, "Systema Naturae". [la icon cite book | last=Linnaeus | first=C | authorlink=Carolus Linnaeus | title=Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. | publisher=Holmiae. (Laurentii Salvii). | date=1758] It is a member of the family Cebidae, the family of New World monkeys containing capuchin monkeys, squirrel monkey, tamarins and marmosets. It is the type species for the genus "Cebus", the genus that includes all the capuchin monkeys.MSW3 Groves|pages=Cebus|id=12100255] It is a member of the "C. capucinus" species group within the genus "Cebus", a group that also includes the White-fronted Capuchin, the Weeper Capuchin and the Kaapori Capuchin (the other species group is the "C. apella" group containing the Tufted Capuchin and others).

Some authorities believe that there are three subspecies of White-headed Capuchin, based on small differences in appearance:
* "C. c. capucinus", from the southern part of the range in Ecuador, Colombia and eastern Panama
* "C. c. imitator", from most of Nicaragua, Costa Rica and western Panama
* "C. c. limitaneus", from Honduras and northern Nicaragua

However, other authorities do not recognize any separate subspecies, and regard "C. c. imitator" and "C. c. limitaneus" as synonyms of "C. capucinus".

References

External links

* [http://www.primatesofpanama.org/academicresources/articles/capuchin.htm Natural history of the White-throated Capuchin]
* [http://www3.interscience.wiley.com/cgi-bin/abstract/110517704/ABSTRACT?CRETRY=1&SRETRY=0 Use of a club by a wild white-faced capuchin to attack a venomous snake]