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Overview

Brief Summary

The capuchin monkeys, formerly all placed in the genus Cebus, are now often separated into two genera, Cebus (for the gracile or untufted capuchins) and Sapajus (for the robust or tufted capuchins). Although the distinction between these two groups was well established by the mid-20th century, the delineation of species and subspecies has been less stable.  Until the early 21st century, however, four species of capuchins were generally recognized: White-faced Capuchin (C. capucinus), White-fronted Capuchin (C. albifrons), the Weeper Capuchin (C. olivaceus or nigrovittatus), and a single robust species, the Brown Capuchin (C. apella). Within these species, various subspecies were often recognized. Based on more recent work, however, including molecular phylogenetic studies, Rylands and Mittermeier (2013) recognized 14 species of Cebus (and eight Sapajus), with the clear understanding that this is unlikely to be the last word on capuchin taxonomy as additional samples and data are acquired and analyzed. They recognized two species of white-faced capuchins, the Colombian White-faced Capuchin (C. capucinus), occurring from northwestern Ecuador to eastern Panama, and the Panamanian White-faced Capuchin (C. imitator), ocurring from western Panama to Honduras (and possibly Belize). See Rylands and Mittermeier (2013) for a review of capuchin biology and systematics and references to key literature.

  • Alfaro, J.W.L., J. De Sousa E. Silva, Jr., and A.B. Rylands. 2012. How Different Are Robust and Gracile Capuchin Monkeys? An Argument for the Use of Sapajus and Cebus. American Journal of Primatology 74: 273-286.
  • Boubli, J.P., A.B. Rylands, I.P. Farias, et al. 2012. Cebus Phylogenetic Relationships: A Preliminary Reassessment of the Diversity of the Untufted Capuchin Monkeys. American Journal of Primatology 74: 381-393.
  • Ruiz-Garcia, M., M.I. Castillo, A. Ledezma, et al. 2012. Molecular Systematics and Phylogeography of Cebus capucinus (Cebidae, Primates) in Colombia and Costa Rica by Means of the Mitochondrial COII Gene. American Journal of Primatology 74: 366-380.
  • Rylands, A.B. and R. Mittermeier. 2013. Family Cebidae (squirrel monkeys and capuchins). Pp. 348-413. In: Mittermeier, R.A., A.B. Rylands, and D.E. Wilson. Handbook of the Mammals of the World,. Volume 3. Primates. Lynx Edicions, Barcelona.
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Distribution

Range Description

Cebus capucinus is the only capuchin monkey in Central America, ranging from Honduras in the north, through Nicaragua, Costa Rica and Panama and through the Chocó-Darién into Colombia (Hernández-Camacho and Cooper 1976; Rodríguez-Luna et al. 1996; Reid, 1997; Marineros and Gallegos 1998). Four subspecies are recognized here:

Cebus capucinus capucinus
In Colombia, the White-throated Capuchin occurs south from the Panamanian border along the Pacific Coast, west of the Andes (up to 1,800–2,000 m above sea level) into northwestern Ecuador. It is restricted to the west bank of the upper Río Cauca (between the Cordillera Occidental and Cordillera Central) and extends north across the Río Sinu into Cordoba, Sucre and Atlantico to the town of Barranquilla on the northern coast of Colombia (Hernández-Camacho and Cooper 1976; Defler 2004). In Central America, C. c. capucinus extends west as far the Panama Canal (Baldwin and Baldwin 1977; Hall 1981).

Cebus capucinus limitaneus
In Honduras, Marineros and Gallegos (1998) recorded this subspecies from throughout the north (Departments of Gracias a Dios, Colón, Atlantida, and Cortés) besides Santa Bárbara in the north-west, and Olancho and El Paraíso in the east. Hollister (1914) also listed a skin from British Honduras (Belize). There have been unauthenticated reports of capuchins in the Mayan Mountains of western Belize (the Chiquebul forest and in the region of the Trio and Bladen branches of the Monkey River) and in Sarstoon National Park on the southern border. Its occurrence in Belize has never been confirmed (McCarthy 1982; Dahl 1984, 1987; Hubrecht 1986). Silva-López et al. (1995; Silva-López 1998) also reported on the possible occurrence of C. c. limitaneus in Guatemala, in the Sierra del Espíritu Santo near the Guatemala-Honduras border. This also remains to be substantiated.

Cebus capucinus imitator
Hall (1981) places this subspecies in western Panama, west from the Canal, and in adjacent areas of Costa Rica. Populations also occur on the islands of Coiba and nearby Jicarón. Baldwin and Baldwin (1976, 1977) documented the occurrence of C. capucinus in a number of localities in the Province of Chiriquí, south-western Panama. Crockett et al. (1997) listed localities in Nicaragua, and Allen (1908, 1910) recorded specimens of Cebus capucinus (referred to as Cebus hypoleucus in Allen, 1908), from Ocotal (northern highlands, 4,500 ft), and localities on the east slope of the highlands, Savala (800 ft), Chontales (lowlands east of Lake Nicaragua, altitudes 500-1,500 ft), and the Río Tuma (500 ft) and Muy Muy (Matagalpa Province, 1,500-2,000 ft).

Cebus capucinus curtus
Gorgona Island, Colombia, possibly introduced in the 16th or 17th centuries.
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Geographic Range

Cebus capucinus is native to Central America in the Neotropical Region. They are found in Honduras, Nicaragua, Costa Rica, Panama, and along the coast of Columbia and Ecuador. Some individuals have been reported as far south as Argentina. This species has one of the widest ranges of all New World monkeys.

Biogeographic Regions: neotropical (Native )

  • Richard, A. 1985. Primates In Nature. United States: W. H. Freeman and Company.
  • Miller, L. 2002. Eat or be Eaten: Predator Sensitive Foraging Among Primates. United Kingdom: Cambridge University Press.
  • Smuts, B., D. Cheney, R. Seyfarth, R. Wrangham, T. Struhsaker. 1987. Primate Societies. United States: The University of Chicago Press.
  • Moynihan, M. 1976. The New World Primates: adaptive radiation and the evolution of social behavior, languages, and intelligence. Princeton, NJ: Princeton University Press.
  • Wolfheim, J. 1983. Primates of the world: distribution, abundance, and conservation. Seattle, Washington: University of Washington Press.
  • Grzimek, B., K. Gold. 2004. New World monkeys I: Squirrel monkeys and capuchins. Pp. 101-113 in Grzimek's Animal Life Encyclopedia, Vol. 14, 2nd Edition. Farmington Hills, Mi: Gale.
  • Estrada, A., P. Garber, M. Pavelka, L. Luecke. 2006. New perspectives in the study of Mesoamerican primates : distribution, ecology, behavior, and conservation. New York: Springer.
  • Kinzey, W. 1997. New World primates : ecology, evolution, and behavior. New York: Aldine de Gruyter.
  • Perry, S. 2003. In The Biology of Traditions: Models and Evidence. Cambridge: University of Camrbridge Press. Accessed September 13, 2008 at http://www.eva.mpg.de/phylogen/pdf/chapter.pdf.
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Physical Description

Morphology

Physical Description

White-faced capuchins have distinctive markings that distinguish them from other capuchin monkeys. Their dorsum and hindquarters are solid black, while the upper chest, forearms, and the fur around the face are white. They have tan-colored facial skin and a black cap on their heads.  White-faced capuchins have prehensile tails, used for grasping and carrying food and for added postural support. They reach their full adult size by age 8. Males weigh between 3 and 4 kg and females weigh between 2 and 3 kg. This is the greatest degree of sexual dimorphism among the capuchins. They are primarily quadrupedal, but are also excellent leapers and climbers.

Range mass: 2 to 4 kg.

Range length: 435 (high) mm.

Other Physical Features: endothermic ; homoiothermic; bilateral symmetry

Sexual Dimorphism: male larger

  • Gerber, P., J. Rehg. 1999. The Ecological Role of the Prehensile Tail in White-Faced Capuchins (Cebus capucinus). American Journal of Physical Anthropology, Volume: 110 Issue: 3: 325-339.
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Type Information

Type for Cebus capucinus
Catalog Number: USNM 16084
Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Mammals
Sex/Stage: Male; Adult
Preparation: Skin; Skull
Collector(s): C. Townsend
Year Collected: 1887
Locality: Segovia River [= Rio Coco], Cabo Gracias a Dios, mouth of Rio Segovia, border with Nicaragua (see Hershkovitz 1949:347), Gracias a Dios, Honduras, North America
  • Type: Hollister, N. 1914 May 11. Proceedings of the Biological Society of Washington. 27: 105.
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Ecology

Habitat

Rio Negro-Rio San Sun Mangroves Habitat

This taxon occurs in the Rio Negro-Rio San Sun mangroves, which consists of a disjunctive coastal ecoregion in parts of Costa Rica, extending to the north slightly into Nicaragua and south marginally into Panama. Furthermore, this species is not necessarily restricted to this ecoregion. Mangroves are sparse in this ecoregion, and are chiefly found in estuarine lagoons and small patches at river mouths growing in association with certain freshwater palm species such as the Yolillo Palm (Raphia taedigera), which taxon has some saline soil tolerance, and is deemed a basic element of the mangrove forest here. These mangrove communities are also part of a mosaic of several habitats that include mixed rainforest, wooded swamps, coastal wetlands, estuarine lagoons, sand backshores and beaches, sea-grasses, and coral reefs.

The paucity of mangroves here is a result of the robust influx of freshwater to the coastline ocean zone of this ecoregion. Among the highest rates of rainfall in the world, this ecoregion receives over six metres (m) a year at the Nicaragua/ Costa Rica national border. Peak rainfall occurs in the warmest months, usually between May and September. A relatively dry season occurs from January to April, which months coincides with stronger tradewinds. Tides are semi-diurnal and have a range of less than one half metre.

Mangroves play an important role in trapping sediments from land that are detrimental to the development of both coral reefs and sea grasses that are associated with them. Mangrove species including Rhizopora mangle, Avicennia germinans, Laguncularia racemosa, Conocarpus erecta and R. harrisonii grow alone the salinity gradient in appropriate areas. Uncommon occurrences of Pelliciera rhizophorae and other plant species associated with mangroves include Leather ferns Acrostichum spp., which also invade cut-over mangrove stands and provide some protection against erosion. In this particular ecoregion, the mangroves are associated with the indicator species, freshwater palm, Raphia taedigera. Other mangrove associated species are Guiana-chestnut ( Pachira aquatica) and Dragonsblood Tree (Pterocarpus officinalis).

Reptiles include the Basilisk Lizard (Basiliscus basiliscus), Caiman (Caiman crocodilus), Green Sea Turtle (Chelonia mydas), Leatherback Turtle (Dermochelys coriacea) and Green Iguana (Iguana iguana). The beaches along the coast within this ecoregion near Tortuguero are some of the most important for nesting green turtles. The offshore seagrass beds, which are among the most extensive in the world, are a source of food and refuge for the endangered Green Sea Turtle (Chelonia mydas). Several species of frogs  of the family Dendrobatidae are found in this mangrove ecoregion as well other anuran species and some endemic salamander taxa.

Mammal species found in this highly diverse ecoregion include: Lowland Paca (Agouti paca), primates such as Mantled Howler Monkey (Alouatta palliata), Geoffrey's Spider Monkey (Ateles geoffroyi), White-faced Capuchin (Cebus capucinus), Brown-throated Sloth (Bradypus variegatus), Silky Anteater (Cyclopes didactylus) and Nine-banded Armadillo (Dasypus novemcintus).  Also found in this ecoregion are carnivores such as Ocelot (Leopardus pardalis),  Central American Otter (Lutra annectens), Jaguar (Panthera onca), Northern Racooon (Procyoon lotor), and Crab-eating Racoon (P. cancrivorus).

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Moist Pacific Coast Mangroves Habitat

This taxon occurs in the Moist Pacific Coast mangroves, an ecoregion along the Pacific coast of Costa Rica with a considerable number of embayments that provide shelter from wind and waves, thus favouring mangrove establishment. Tidal fluctuations also directly influence the mangrove ecosystem health in this zone. The Moist Pacific Coast mangroves ecoregion has a mean tidal amplitude of three and one half metres,

Many of the streams and rivers, which help create this mangrove ecoregion, flow down from the Talamanca Mountain Range. Because of the resulting high mountain sediment loading, coral reefs are sparse along the Pacific coastal zone of Central America, and thus reef zones are chiefly found offshore near islands. In this region, coral reefs are associated with the mangroves at the Isla del Caño Biological Reserve, seventeen kilometres from the mainland coast near the Térraba-Sierpe Mangrove Reserve. The Térraba-Sierpe, found at the mouths of the Térraba and Sierpe Rivers, is considered a wetland of international importance.

Because of high moisture availability, the salinity gradient is more moderate than in the more northern ecoregion such as the Southern dry Pacific Coast ecoregion. Resulting mangrove vegetation is mixed with that of marshland species such as Dragonsblood Tree (Pterocarpus officinalis), Campnosperma panamensis, Guinea Bactris (Bactris guineensis), and is adjacent to Yolillo Palm (Raphia taedigera) swamp forest, which provides shelter for White-tailed Deer (Odocoileus virginianus) and Mantled Howler Monkeys (Alouatta palliata). Mangrove tree and shrub taxa include Red Mangrove (Rhizophora mangle), Mangle Caballero (R. harrisonii) R. racemosa (up to 45 metres in canopy height), Black Mangrove (Avicennia germinans) and Mangle Salado (A. bicolor), a mangrove tree restricted to the Pacific coastline of Mesoamerica.

Two endemic birds listed by IUCN as threatened in conservation status are found in the mangroves of this ecoregion, one being the Mangrove Hummingbird (Amazilia boucardi EN), whose favourite flower is the Tea Mangrove (Pelliciera rhizophorae), the sole mangrove plant pollinated by a vertebrate. Another endemic avain species to the ecoregion is the  Yellow-billed Cotinga (Carpodectes antoniae EN).  Other birds clearly associated with the mangrove habitat include Roseate Spoonbill (Ajaia ajaja), Gray-necked Wood Rail (Aramides cajanea), Rufous-necked Wood Rail (A. axillaris), Mangrove Black-hawk (Buteogallus anthracinus subtilis),Striated Heron (Butorides striata), Muscovy Duck (Cairina moschata), Boat-billed Heron (Cochlearius cochlearius), American White Ibis (Eudocimus albus), Amazon Kingfisher (Chloroceryle amazona), Mangrove Cuckoo (Coccyzus minor), Yellow Warbler (Setophaga petechia), and Black-necked Stilt (Himantopus mexicanus VU) among other avian taxa.

Mammals although not as numerous as birds, include species such as the Lowland Paca (Agouti paca), Mantled Howler Monkey (Alouatta palliata), White-throated Capuchin (Cebus capucinus), Silky Anteater (Cyclopes didactylus), Central American Otter (Lontra longicaudis annectens), White-tailed Deer (Odocoileus virginianus), feeds on leaves within A. bicolor and L. racemosa forests. Two raccoons: Northern Raccoon (Procyon lotor) and Crab-eating Raccoon (P. cancrivorus) can be found, both on the ground and in the canopy consuming crabs and mollusks. The Mexican Collared Anteater (Tamandua mexicana) is also found in the Moist Pacific Coast mangroves.

There are a number of amphibians in the ecoregion, including the anuran taxa: Almirante Robber Frog (Craugastor talamancae); Chiriqui Glass Frog (Cochranella pulverata); Forrer's Grass Frog (Lithobates forreri), who is found along the Pacific versant, and is at the southern limit of its range in this ecoregion. Example salamanders found in the ecoregion are the Colombian Worm Salamander (Oedipina parvipes) and the Gamboa Worm Salamander (Oedipina complex), a lowland organism that is found in the northern end of its range in the ecoregion. Reptiles including the Common Basilisk Lizard (Basiliscus basiliscus), Boa Constrictor (Boa constrictor), American Crocodile (Crocodilus acutus), Spectacled Caiman (Caiman crocodilus), Black Spiny-tailed Iguana (Ctenosaura similis) and Common Green Iguana (Iguana iguana) thrive in this mangrove ecoregion.

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Habitat and Ecology

Habitat and Ecology
In Central America, present in dry deciduous forests on Pacific coast (rainfall less than 1,750 mm) and, humid subtropical forests on the Caribbean coast (rainfall as much as 5,000 mm), mangroves and second growth (Freese and Oppenheimer 1981: Reid 1997). In Colombia, C. capucinus occurs in primary and advanced secondary forest remnants, degraded forests and forests with large concentrations of palms, especially Scheelea magdalenica at altitudes up to 2,000 m above sea level (Defler 2004). In Ecuador, it occurs in humid tropical and sutropical forests from sea level to 1,800 m (Tirira 2007).

Capucins are frugivores-insectivores, including wide variety of fruits, seeds and arthropods, frogs, nestlings and even small mammals, supplemented by stems, flowers and leaves. They are extractive, manipulative foragers. Males disperse. Both sexes take up linear hierarchies, the top ranking male being dominant to the top ranking female. Multi-male groups range in size from 5-30 individuals. Fedigan and Jack (2001) recorded a mean group size of 15.8 for 20 groups in the Santa Rosa National Park, Costa Rica. The adult sex-ratio was 0.71. In Chiriqui, Panama, Baldwin and Baldwin (1977) found C. capucinus occupying small forest patches of 0.2 to 40 ha in small groups of 2-5 individuals, whereas groups in larger forests reached sizes do 20 or more iindividuals.

Although new born offspring can be seen year round, C. capucinus in the Santa Rosa National Park, Costa Rica, shows a birth peak in the dry season and early rainny season (Fedigan et al. 1996).

Size:
HB 33.0-49.5 cm, TL 39.7-54.0 cm.
Weight 1.8-4.3 kg (Reid 1997).

Systems
  • Terrestrial
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White-faced capuchins are highly adaptable and occupy a wide range of habitats. They live in a variety of wet, dry, primary, and secondary forests, but prefer tropical evergreens and dry deciduous forests. White-faced capuchins have a preference for close-canopied forests up to as high as 2100 m but mainly occupy the middle strata around 1100 m. They are common in areas high in humidity and well-drained lowlands. Occasionally, they have been found in volcanic foothills and coastal plains.

Range elevation: 0 to 2100 m.

Average elevation: 1100 m.

Habitat Regions: tropical ; terrestrial

Terrestrial Biomes: forest ; rainforest

Other Habitat Features: riparian

  • Eisenberg, J. 1989. Mammals of the Neotropics. Chicago: University of Chicago Press.
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Trophic Strategy

Food Habits

White-faced capuchins eat a wide variety of plants and animals. Their primary diet is a variety of fruits and nuts, but insects, other invertebrates, and small vertebrates are taken opportunistically. Vertebrates eaten include squirrels, tree rats, lizards, and birds. The diet varies regionally and seasonally, but generally consists of about 50 to 80% fruit, 20 to 30% animal material, and 10% other plant material. White-faced capuchins eat frequently and are adventurous in their food choices. They will try almost anything once and learn through trial and error about what is edible or desirable. One study showed that they ate 63 different plant species from 34 families at Santa Rosa Park. They are excellent foragers from a very young age. As young as 1 year old they are able to seek out food almost as well as adults, their only limitations being size and strength.

Animal Foods: birds; mammals; amphibians; reptiles; insects; terrestrial non-insect arthropods

Plant Foods: seeds, grains, and nuts; fruit; flowers; sap or other plant fluids

Primary Diet: herbivore (Frugivore ); omnivore

  • Chapman, C., L. Fedigan. 1990. Dietary Differences between Neighboring Cebus Capucinus Groups: Local traditions, Food availability or responses to food profitability. Folia Primatologica, 54/3-4: 177-186.
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Associations

Ecosystem Roles

White-faced capuchins play an important role in seed dispersal, influencing forest regeneration.  Blood-born nematode (roundworm) parasites of this monkey include: Microfilaria panamensis, Tetrapetalonema panamensis, and Dipetalonema obtusa.

Ecosystem Impact: disperses seeds

Commensal/Parasitic Species:

  • Microfilaria panamensis
  • Tetrapetalonema panamensis
  • Dipetalonema obtusa

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Predation

The most common predators of white-face capuchins are snakes, especially tree boas (Corallus) and lanceheads (Bothrops). Caimans, cats, such as jaguars and ocelots and large raptors, such as harpy eagles, also prey on these capuchins. White-face capuchins sound alarm calls when they spot a predator. Living in tight-knit social groups helps them to stay vigilant against predators. Groups threatened by a predator will either flee or mob the predator.

Known Predators:

Anti-predator Adaptations: cryptic

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Life History and Behavior

Behavior

Communication and Perception

White-faced capuchins are highly social and communication is very important. Individuals spend a great deal of time in social bonding and establishing trust. And example is hand-sniffing, where one monkey will stick his/her fingers in the other monkey's nose and then other monkey repeats the activity. This can last for several minutes and is done with a trance-like expression. They may also suck on each other's fingers and tails for long periods of time. White-faced capuchins also play mouth games, where one individual will insert something into their mouth, whether it be the other monkey's finger, a patch of fur, or some inanimate object. The object of this game is to pry the item out of the others mouth, and then the object is either placed back in the mouth to start a new game, or they take turns and switch roles. When predators approach, white-faced capuchins use trill vocalizations to coordinate movement in the group. A different alarm call is used to alert others that a predator or intruder in nearby.

Communication Channels: visual ; tactile ; acoustic ; chemical

Other Communication Modes: mimicry

Perception Channels: visual ; acoustic

  • Radick, G. 2007. The Simian Tongue: The Long Debate about Animal Language. Chicago and London: The University of Chicago Press.
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Life Expectancy

Lifespan/Longevity

White-faced capuchin maximum lifespan in captivity is 55 years. Wild lifespan may be less than half of that. Factors such as predation, disease, and infection from wounds sustained while fighting are all responsible for many deaths each year. Tree removal, logging, and clearcutting are indirectly the leading cause of white-face capuchin death by drastically reducing suitable habitats.

Range lifespan

Status: captivity:
55 (high) years.

Typical lifespan

Status: wild:
30 (high) years.

Typical lifespan

Status: captivity:
45 (high) years.

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Lifespan, longevity, and ageing

Maximum longevity: 54 years (captivity) Observations: One wild-born male called "Bobo" lived 53.5 years in captivity, making it at least 54 years of age when he died at Lederle Laboratories. While this is the only record of a specimen of this genus to live more than 50 years, another wild-born male specimen lived 46.9 years at Mesker Park Zoo, making it about 48 years-old when he died and making it the second longest-lived specimen of the genus (Richard Weigl 2005).
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Reproduction

White-faced capuchins are polygamous, with males and females having multiple partners. There is a dominance hierarchy among males, with alpha males having more mating opportunities. However, subordinate males are also allowed to mate. Alpha males spend a large portion of their time engaged in deterring predators and males from outside groups. Providing his group with this protection gives the alpha male precedence over the other males in mating opportunities. Males and females engage in a set of specific vocalizations, facial expressions, and postures before copulation.

Mating System: polygynandrous (promiscuous)

Females give birth to a single offspring at about 2 year intervals. Breeding is seasonal, with peak female fertility from January to April. Mating occurs outside of these months but rarely results in conceptions. Gestation last for an average of 160 days. Adult females become sexually mature around age 4, but typically don't give birth until age 7. Males become sexually mature at 7 to 10 years old.

Breeding interval: White-faced capuchins breed year-round, though females only conceive once every 2 years, on average.

Breeding season: Peak breeding is between January and April.

Range number of offspring: 1 (high) .

Average number of offspring: 1.

Range gestation period: 157 to 167 days.

Average gestation period: 160 days.

Average birth mass: 230 g.

Range weaning age: 2 to 4 months.

Range time to independence: 4 to 8 years.

Average time to independence: 7 years.

Range age at sexual or reproductive maturity (female): 4 to 7 years.

Average age at sexual or reproductive maturity (female): 5 years.

Range age at sexual or reproductive maturity (male): 7 to 10 years.

Average age at sexual or reproductive maturity (male): 8 years.

Key Reproductive Features: iteroparous ; seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); fertilization ; viviparous

Primary care for infants is provided by the mother. Females carry, protect, and feed the young until they are able to take care of themselves. Males do not help with care of infants, but may assist young in the social hierarchy once they are independent. Alpha males help to protect members of their group from intruders and predators.

Parental Investment: altricial ; pre-fertilization (Provisioning, Protecting: Female); pre-hatching/birth (Provisioning: Female, Protecting: Female); pre-weaning/fledging (Provisioning: Female, Protecting: Male, Female); pre-independence (Provisioning: Female, Protecting: Male, Female); post-independence association with parents; extended period of juvenile learning

  • Eisenberg, J. 1989. Mammals of the Neotropics. Chicago: University of Chicago Press.
  • Smuts, B., D. Cheney, R. Seyfarth, R. Wrangham, T. Struhsaker. 1987. Primate Societies. United States: The University of Chicago Press.
  • Grzimek, B., K. Gold. 2004. New World monkeys I: Squirrel monkeys and capuchins. Pp. 101-113 in Grzimek's Animal Life Encyclopedia, Vol. 14, 2nd Edition. Farmington Hills, Mi: Gale.
  • Estrada, A., P. Garber, M. Pavelka, L. Luecke. 2006. New perspectives in the study of Mesoamerican primates : distribution, ecology, behavior, and conservation. New York: Springer.
  • Kinzey, W. 1997. New World primates : ecology, evolution, and behavior. New York: Aldine de Gruyter.
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Conservation

Conservation Status

IUCN Red List Assessment


Red List Category
LC
Least Concern

Red List Criteria

Version
3.1

Year Assessed
2008

Assessor/s
Causado, J., Cuarón, A.D., Shedden, A., Rodríguez-Luna, E. & de Grammont, P.C.

Reviewer/s
Mittermeier, R.A. & Rylands, A.B. (Primate Red List Authority)

Contributor/s

Justification
This species is listed as Least Concern as it is widespread, exhibits an adaptability to a variety of habitats, and there are few major threats driving a significant overall population decline at present.

History
  • 2003
    Least Concern
    (IUCN 2003)
  • 2003
    Least Concern
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White-face capuchins are widespread and their populations are not currently considered threatened. The main pressures they face are habitat degradation, deforestation, and being hunted for food.

US Federal List: no special status

CITES: appendix ii

State of Michigan List: no special status

IUCN Red List of Threatened Species: least concern

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Population

Population
No information on population densities are available for this species.

Population Trend
Unknown
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Threats

Major Threats
Although habitat loss is certainly a threat, all subspecies have a varied diet, and are able to live in a variety of different habitats and disturbed areas. They are sometimes kept as pets and hunted.

There are no current major threats to Cebus capucinus curtus on Gorgona Island, which is a protected area (Parque Nacional Gorgona). However, as this is the only location in which this subspecies is found (Gorgona Island has an area of 15 km², it remains very susceptible to any natural or anthropogenic threats in the near future.
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Management

Conservation Actions

Conservation Actions
This species is listed on CITES Appendix II. It is present in a number of protected areas:

Cebus capucinus capucinus

Panama
Porto Belo National Park (34,848 ha) (Matamoros and Seal 2001)
Chagres National Park (129,000 ha) (Matamoros and Seal 2001)
Darién National Park (555,000 ha) (Matamoros and Seal 2001)

Ecuador
Cotacachi-Cayapas Ecological Reserve (Tirira 2007)
Los Cedros Protected Forest (Tirira 2007)

Cebus capucinus imitator

Costa Rica
Santa Rosa National Park (21,913 ha) (Reid 1997; Matamoros and Seal 2001)
Rincón La Vieja National Park (14,083 ha) (Matamoros and Seal 2001)
Piedras Blancas National Park (14,100 ha) (in range)
Chirripó National Park (50,150 ha) (Matamoros and Seal 2001)
Corcovado National Park (41,788 ha) (in range)
Guanacaste National Park (33,786 ha) (Chapman et al. 1989)
Palo Verde National Park (5,704 ha) (Reid 1997)
Cabo Blanco Strict Nature Reserve (14,258 ha) (Matamoros and Seal 2001)

Panama
La Amistad International Park (207,000 ha) (Matamoros and Seal 2001)
Soberanía National Park (22,104 ha) (Matamoros and Seal 2001)
El Copé – Comar Torrijos Herrera (25,275 ha) (Matamoros and Seal 2001)
Volcán Baru National Park (14,000 ha) (Matamoros and Seal 2001)
Altos de Campaña National Park (4,816 ha) (in range)
Cerro Hoya National Park (32,557 ha) (Matamoros and Seal 2001)
Isla Coiba National Park and Jicarón (270,125 ha) (Matamoros and Seal 2001)
El Montuoso Forest Reserve (10,375 ha) (Matamoros and Seal 2001)
Isla Barro Colorado Natural Monument (5,600 ha) (Matamoros and Seal 2001)

Cebus capucinus limitaneus

Honduras
Montaña de Cusuco National Park (18,000 ha) (Marineros and Gallegos 1998; Matamoros and Seal 2001)
Pico Bonito National Park (68,000 ha) (Marineros and Gallegos, 1998; Matamoros and Seal 2001)
Montaña de Yoro National Park (15,500 ha) (Matamoros and Seal 2001)
Sierra de Agalta National Park (27,000 ha) (Marineros and Gallegos 1998; Matamoros and Seal 2001)
Punto Isopo Wildlife Refuge (11,200 ha) (Marineros and Gallegos 1998)

Cebus capucinus curtus

Colombia
Isla Gorgona National Natural Park (1568 ha)
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Relevance to Humans and Ecosystems

Benefits

Economic Importance for Humans: Negative

White-faced capuchins can be carriers of various diseases while in the wild, and because they are caught for the pet trade, these disease can be transmitted to humans. White-faced capuchins are known carriers of malaria and microfilaria. These diseases appear to be more prevalent in infants and juveniles, possibly because of their weaker immune systems. White-faced capuchins can also eat fruit crops.

Negative Impacts: crop pest; causes or carries domestic animal disease

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Economic Importance for Humans: Positive

White-face capuchins are hunted for food and commonly kept as pets and collected in zoos. Most capuchins in zoos are bred in captivity; few are from the wild.

Positive Impacts: pet trade ; food ; research and education

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Wikipedia

White-headed capuchin

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, the white-headed capuchin is important to rainforest ecology for its 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 paraplegic persons. It is a medium-sized monkey, weighing up to 3.9 kg (8.6 lb). 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, the white-headed capuchin is versatile, living in many different types of forest, and eating many different types of food, including fruit, other plant material, invertebrates, and small vertebrates. It lives in troops that can exceed 20 animals and include both males and females. It is noted for its tool use, including rubbing plants over its body in an apparent use of herbal medicine, and also using tools as weapons and for getting to food. It is a long-lived monkey, with a maximum recorded age of over 54 years.

Taxonomy[edit]

Some authorities consider this a member of the subspecies Cebus capucinus imitator.

The white-headed capuchin was one of the many species originally described by Linnaeus in his 18th century work, Systema Naturae.[4] It is a member of the family Cebidae, the family of New World monkeys containing capuchin monkeys, squirrel monkeys, tamarins and marmosets. It is the type species for the genus Cebus, the genus that includes all the capuchin monkeys.[5] 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.[5]

There is disagreement among primatologists about whether there are any subspecies of white-headed capuchin. Some authorities consider there to be three subspecies of white-headed capuchin, based on small differences in appearance:[3]

However, other authorities do not recognize any separate subspecies, and regard C. c. imitator and C. c. limitaneus as synonyms of C. capucinus.[1]

Physical description[edit]

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.[6][7] The white-headed capuchin has mostly black fur, with white to yellow like fur on the neck, throat, chest, shoulders, and upper arms.[8] The face is pink or a white-cream color and may have identifying marks such as dark brows or dark fur patches.[8][9][10] An area of black fur on the crown of the head is distinctive.[8][11] It has a prehensile tail that is often held coiled, giving the white-headed capuchin the nickname "ringtail".[8][12]

Adults reach a length of between 335 and 453 mm (13.2 and 17.8 in), excluding tail, and a weight of up to 3.9 kg (8.6 lb).[8][11] The tail is longer than the body, at up to 551 mm (21.7 in) in length.[8][11] Males are about 27% larger than females.[13] The brain of a white-headed capuchin is about 79.2 g (2.79 oz), which is larger than that of several larger monkey species, such as the Mantled Howler.[11][14]

Behavior[edit]

Social structure[edit]

Walking on four limbs

The white-headed capuchin is a diurnal and arboreal animal.[8] However, it does come down to the ground more often than many other New World monkeys.[15] It moves primarily by walking on all four limbs.[16] It lives in troops, or groups, of up to 40 monkeys (mean 16, range 4–40)[17] and has a male/female adult sex ratio of 0.71 on average (range 0.54–0.88).[17] With rare exceptions, females spend their entire lives with their female kin.[17][18][19] Males migrate to new social groups multiple times during the course of their lifetimes, migrating for the first time between 20 months and 11 years of age.[20][21] The median age of migration in the Santa Rosa population is 4.5 years.[20] Males sometimes migrate alone, but more often they migrate in the company of other males who are often their kin.[19][20][21] One of the unusual features of the kinship structure of the white-headed capuchin, relative to other primate species, is the high degree of relatedness within groups that results from the long tenures of alpha males who sire most of the offspring.[22][23] Alpha males have been known to keep their positions as long as 17 years in this species[22][23] and this puts them in the unusual position of being available to sire the offspring of their daughters and granddaughters, who produce their first offspring at about 6–7 years of age.[17][23] Typically, however, alpha males do not breed with their own daughters, even though they do sire virtually all offspring produced by females unrelated to them.[22] Those subordinate males who are allies of the alpha male in group defense are the males who sire the offspring of the alpha male's daughters. The high degree to which alpha males monopolize matings results in an unusually large number of paternal half-siblings and full siblings in this species relative to other primate species.[23]

Kinship is an important organizing factor in the structuring of female-female social relationships.[23] Particularly in larger groups, females preferentially associate with, groom, and provide coalitionary support to their matrilineally related female kin. They do not exhibit a similar preference for their paternal half sisters, which may mean that they only are capable of recognizing kinship through the maternal line.[23] Dominance rank is also an important organizing factor, with females more often grooming and associating with females who are closer to them in the dominance hierarchy.[23] Female-female dyads groom far more than male-female and male-male dyads.[24] Coalitionary aggression is common both among males and females, and capuchins seem to have an excellent understanding of the alliance structure in their group. For example, when capuchins are fighting, they sensibly recruit aid from someone who is both higher ranking than they are and also better friends with themselves than with their opponent.[25]

Female capuchins have linear dominance hierarchies.[24][26] In contrast to many Old World monkeys such as macaques, in which females socially inherit the rank just below their mothers and just above their next oldest sisters, capuchins do not have a highly predictable ranking within their matrilines.[23] Males are typically dominant to females.[27] The alpha male is always easy to discern, but there are sometimes ambiguous rankings among subordinate males.[19][28] Male-male relationships are tense, and affiliation between males is typically expressed by resting in contact, playing, or non-conceptive sex rather than by grooming.[28][29] Males cooperate in coalitions against potential predators, and also in defense of the group against other males.[19][28][30] Occasionally male coalitionary aggression becomes so violent that males are killed, particularly if they are encountered roaming the forest unaccompanied by allies.[19][31] Because aggression from other male capuchins is the leading cause of death (aside from poaching by humans, where there is contact between humans and capuchins), male allies are critical for self-defense during migration, and to assist in taking over other groups.[19] Male emigration to a new troop typically occurs about every 4 years, so most males are in constant danger of having to defend themselves against other groups of males.[7][32][32][33]

Immigrating males often kill young infants when they take over a group.[19][34][35] Females band together to defend their infants from infanticidal males, but they rarely succeed in saving their infants.[19] Because infants inhibit their mothers from ovulating by nursing frequently, males are able to bring females into estrus earlier by killing the infants and thereby terminating nursing; this has the effect of increasing their breeding opportunities.[19][35][36] Females do often mate with the killers of their infants, and with time, they typically become as supportive of the new alpha male as they had been of the previous one.[19] The alpha male helps defend females from subordinate males within the group as well as from infanticidal males from other groups.[19][37]

Interactions between groups[edit]

White-faced capuchin troops occupy home ranges of between 32 and 86 hectares (79 and 213 acres).[11] They travel between 1 and 3 kilometres (0.62 and 1.86 mi) daily, averaging 2 kilometres (1.2 mi) per day.[38] Although they engage in activity that has been described as "territorial", more recent research indicates that White-faced capuchin troops tend to behave aggressively to other White-faced capuchin troops regardless of where they meet, and the aggression is not necessarily intended to exclude the other troops from a specific home range.[39]

Home ranges overlap extensively,[40][41] so groups are not territorial in the strictest sense of the word. Perhaps because of the intensity of male-male competition and the threat of infanticide, interactions between groups are typically hostile: the males display aggressively toward one another and sometimes engage in physical aggression (even killing an opponent), while females grab their infants and run.[31][40] Typically, males are the primary participants in aggressive intergroup encounters, and it seems likely that males are defending access to the females in their groups.[40] Alpha males, who have the largest reproductive stake in the group, participate at a higher rate than subordinate males.[40] Groups with more males have an advantage over groups with fewer males, but the location of the encounter within the home range matters as well; smaller groups defeat larger groups when the contest occurs in the core or center area of the smaller group's home range.[41]

Interspecific interactions[edit]

The white-headed capuchin sometimes interacts with other sympatric monkey species. White-headed capuchins sometimes travel with and even groom Geoffroy's Spider Monkeys.[11][38] However, aggressive interactions between the capuchins and spider monkeys also occur.[42] Interactions between the white-headed capuchin and mantled howler are infrequent, and sometimes result in the capuchins threatening the larger howlers.[38] However, affiliative associations between the capuchins and howlers do sometimes occur, mostly involving juveniles playing together.[42]

Although South American capuchin species often travel with and feed together with squirrel monkeys, the white-headed capuchin only rarely associates with the Central American squirrel monkey. This appears to be related to the patchier, more dispersed distribution of food resources in Central America and the fact that there is less dietary overlap between the Central American squirrel monkey and the white-headed capuchin than between their South American counterparts. Therefore, there is less benefit to the Central American squirrel monkey in associating with the white-headed capuchin in order to exploit the capuchin's knowledge of food resource distribution. In addition, compared to their South American counterparts, male white-headed capuchins are relatively more alert to rival males than to predators, reducing the predator detection benefits that the Central American squirrel monkey receives from associating with the white-headed capuchin compared to its South American counterparts. Since the squirrel monkeys generally initiate interactions with the capuchins in South America, the fact that similar associations would impose higher foraging costs and impart fewer predator detection benefits to the Central American squirrel monkey leads to fewer associations with the white-headed capuchin.[13][43][44][45]

Several non-primate animal species tend to follow troops 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.[38] 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.[38]

Diet[edit]

The white-headed capuchin is an omnivore. Its primary foods are fruit and insects.[7] It forages at all levels of the forest, including the ground.[38] Methods for finding food include stripping bark off of trees, searching through leaf litter, breaking dead tree branches, rolling over rocks, and using stones as anvils to crack hard fruits.[46] Its prehensile tail assists with feeding, helping support the monkey when foraging for food below the branches.[38]

Foraging in the trees

Fruit can make up between 50% and 67% or more of the capuchin's diet.[7] In one study in Panama, white-headed capuchins ate 95 different fruit species.[7] Among its 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.[47] It generally only eats ripe fruit, testing for ripeness by smelling, tasting and prodding the fruit.[7] It typically eats only the pulp and juice, spitting out the seeds and fibers.[7] Other plant matter eaten includes flowers, young leaves, seeds of certain plants, and bromeliads.[7][48] It also uses the bromelids as a water source, drinking the water that gets trapped inside.[7] In Carara National Park the capuchins have a varied diet in addition to the above of banana fruits and flowers, heliconia seeds, huevos de caballo fruits and anacardiaceae stems.[49]

Insect prey eaten includes beetle larvae, butterfly and moth caterpillars, ants, wasps, and ant and wasp larvae.[7] It also eats larger prey, such as birds, bird eggs, frogs, lizards, crabs, mollusks and small mammals.[7][50] The population in Guanacaste, Costa Rica in particular is noted for hunting squirrels, magpies, White-crowned Parrots[7] and baby coatis.[51] The amount of vertebrate prey eaten varies by troop.[7] Even neighboring troops can show significant differences in their diets.[46]

The diet can vary between the rainy and dry season. For example, in Guanacaste, Costa Rica the white-headed capuchin can eat a wide variety of fruits as well as caterpillars in the early rainy season (June to November).[47] But during the dry season, only figs and a few other types of fruit are available.[47] During the dry season, chitinous insects, ant and wasp larvae and vertebrates become a particularly important part of the white-headed capuchin's diet.[47] Access to water can also become an issue during the dry season. The white-headed capuchin likes to drink daily, so in forests where water holes dry up during the dry season, there can be competition between troops over access to the remaining water holes.[47]

Tool use[edit]

Capuchins are considered among the most intelligent of the New World monkeys; they have been the subject of many studies on behaviour and intelligence. The capuchins' intelligence is thought to be an adaptation to support 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.[52]

The white-headed capuchin is known to rub parts of certain plants into their hair. Plants used in this manner include citrus fruits, vines of the genera Piper and Clematis, monkey comb (genus Sloanea), dumb cane and custard apple.[7][50] Ants and millipedes are also used in this way.[7] It is not definitively known what this 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.[7] Alternatively, it may be a form of scent marking.[7]

The white-headed capuchin also uses tools in other ways. It has been known to beat snakes with sticks in order to protect itself or to get the snake to release an infant.[7] In captivity, it has been known to use tools to get to food or to defend itself, and in one case a white-headed capuchin used a squirrel monkey as a projectile, hurling it at a human observer.[7] It has been historically noted that the species is often able to recognize, and therefore avoid baited cage traps, and hidden net snares are often the only way to capture this monkey.[citation needed] Some populations also use trees or other hard surfaces as anvils in order to crack mollusks.[50] And it sometimes uses sticks as probes to explore openings.[53]

Though the white-headed capuchin has perhaps the most extensive and most frequent tool use in comparison to the other gracile capuchins, its tool use is considerably inferior to that of robust capuchins, especially the tufted capuchin.[citation needed] Factors such as easier access to water and food may have to do with the white-headed capuchin's less extensive use of tools.[citation needed]

The white-headed capuchin's intelligence and ability to use tools allows them to be trained to assist paraplegics.[54] Other species of capuchin monkeys are also trained in this manner.[55] White-headed capuchins can also be trained for roles on television and movies, such as Marcel on the television series Friends.[56] They were also traditionally used as organ grinder monkeys.[57]

Communication[edit]

Facial expression

The white-headed capuchin is noisy.[8] Loud calls, such as barks and coughs, are used to communicate threat warnings, and softer calls, such as squeals, are used in intimate discourse.[7] Different types of threats, such as a threat from a terrestrial animal versus a threat from a bird, invoke different vocalizations.[38] Facial expressions and scent are also important to communication.[58] It sometimes engages in a practice known as "urine washing", in which the monkey rubs urine on its feet.[59] The exact purpose of this practice is unknown, but it may be a form of olfactory signal.[58]

Reproduction[edit]

The white-headed capuchin uses a polygamous mating system in which a male may mate with multiple females.[38] Although the dominant male does not monopolize breeding, studies have shown that the dominant male does tend to father most of the young.[33] Although a female may mate with several males, the dominant male may be more likely to copulate when the female is at peak fertility.[33][60] Nonetheless, there is evidence that dominant males do tend to avoid breeding with their own daughters who are members of the troop.[61] Such avoidance is rare among New World primates.[61]

Copulation takes about 2 minutes, and the gestation period is 5 to 6 months.[38] Usually a single young is born, but twins occur occasionally. Most births occur during the dry season from December to April.[11][38] The infant is carried across its mother's back for about 6 weeks.[38] After about 4 to 5 weeks it can stray from its mother for brief periods and by about 3 months it can move around independently, although some infants will be mostly independent earlier. Weaning occurs between 6 and 12 months. While the mother rests, the young spends most of its time foraging or playing, either on its own or with other juveniles.[38] Capuchins engage in high levels of alloparenting, in which monkeys other than the mother help care for the infant.[62] Infants are carried by alloparents most often between 4 and 6 weeks in age.[19] Males as well as females engage in alloparenting.[19][38]

Like other capuchin species, the white-headed capuchin matures slowly. Sexual maturity can be reached at 3 years.[57] But on average, females give birth for the first time at 7 years old and give birth every 26 months thereafter.[13] Males attain reproductive maturity at 10 years old.[13] The white-headed capuchin has a long life span given its size. The maximum recorded life span in captivity is over 54 years.[13]

Distribution and habitat[edit]

White-headed capuchin at Frío River, 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.[3] It has also been reported to occur in eastern Guatemala and southern Belize, but these reports are unconfirmed.[3] 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.[3] It 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.[63]

It should also be noted that while it is very common in Costa Rica and Panama, the monkey has been largely extirpated from Honduras and much of Nicaragua. Many Honduran Capuchins were captured and relocated to the island of Roatán, and many Nicaraguan Capuchins were captured and relocated to the island of Ometepe. In Nicaragua, wild Capuchins may still be easily spotted in regions around Masaya, as well as around Bluefields and other locations around the South Caribbean coast. The monkey can also be observed near the Costa Rican border along the San Juan River, and in Kahka Creek Rainforest Preserve.[citation needed]

It is 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.[8][57] However, it appears to prefer primary or advanced secondary forests.[38] Also, higher densities of white-headed capuchins are found in older areas of forest and in areas containing evergreen forest, as well as areas with more water availability during the dry season.[64]

Conservation status[edit]

The white-headed capuchin is regarded as "least concern" from a conservation standpoint by IUCN. However, its numbers are impacted by the fact that it is sometimes captured for the pet trade.[7] Its status can also be harmed by deforestation. However, deforestation may also impact its 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.[7] The white-headed capuchin can adapt to forest fragmentation better than other species due to its ability to live in a wide variety of forest types and exploit a wide variety of food sources.[65] The white-headed capuchin is important to its ecosystems as a seed and pollen disperser.[7][57] It also impacts the ecosystem by eating insects that act as pests to certain trees, by pruning certain trees, such as Gustavia superba and Bursera simaruba, causing them generate more branches and possibly additional fruit, and by accelerating germination of certain seeds when they pass through the capuchin's digestive tract.[7] In addition, the white-headed capuchin sometimes kills Acacia collinsii plants when it rips through the plant's branches to get to resident ant colonies.[7]

References[edit]

  1. ^ a b Groves, C. P. (2005). Wilson, D. E.; Reeder, D. M, eds. Mammal Species of the World (3rd ed.). Baltimore: Johns Hopkins University Press. p. 137. OCLC 62265494. ISBN 0-801-88221-4. 
  2. ^ Causado, J., Cuarón, A.D., Shedden, A., Rodríguez-Luna, E. & de Grammont, P.C. (2008). "Cebus capucinus". IUCN Red List of Threatened Species. Version 2011.2. International Union for Conservation of Nature. Retrieved 19 January 2012. 
  3. ^ a b c d e Rylands, A., Groves, C., Mittermeier, R., Cortes-Ortiz, L., and Hines, J. (2006). "Taxonomy and Distributions of Mesoamerican Primates". In Estrada, A., Garber, P., Pavelka, M. & Luecke, L. New Perspectives in the Study of Mesoamerican Primates. New York: Springer. pp. 40–43. ISBN 0-387-25854-X. 
  4. ^ (Latin) Linnaeus, C (1758). Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata (10 ed.). Holmiae. (Laurentii Salvii). p. 29. 
  5. ^ a b Groves, C. P. (2005). Wilson, D. E.; Reeder, D. M, eds. Mammal Species of the World (3rd ed.). Baltimore: Johns Hopkins University Press. p. Cebus. OCLC 62265494. ISBN 0-801-88221-4. 
  6. ^ "Capuchin Franciscans F.A.Q.". Capuchin Franciscans Vocation Office Province of Saint Joseph. Retrieved 2008-09-01. 
  7. ^ a b c d e f g h i j k l m n o p q r s t u v w x y Wainwright, M. (2002). The Natural History of Costa Rican Mammals. Zona Tropical. pp. 135–139. ISBN 0-9705678-1-2. 
  8. ^ a b c d e f g h i Emmons, L. (1997). Neotropical Rainforest Mammals A Field Guide (Second ed.). University of Chicago Press. pp. 130–131. ISBN 0-226-20721-8. 
  9. ^ Luedtke, Karen (2012). Jungle Living: A look at life and social behavior of man and monkey in Central American. pp. 40–45. ISBN 0-9832448-2-0. 
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