White-headed Capuchin
Overview
The white-headed capuchin (Cebus capucinus), also known as the white-faced capuchin or white-throated capuchin, is a medium-sized New World monkey
New World monkey
New World monkeys are the five families of primates that are found in Central and South America: Callitrichidae, Cebidae, Aotidae, Pitheciidae, and Atelidae. The five families are ranked together as the Platyrrhini parvorder and the Ceboidea superfamily, which are essentially synonymous since...

 of the family Cebidae
Cebidae
The Cebidae is one of the five families of New World monkeys now recognised. It includes the capuchin monkeys and squirrel monkeys. These species are found throughout tropical and subtropical South and Central America.-Characteristics:...

, subfamily Cebinae. Native to the forests of Central America
Central America
Central America is the central geographic region of the Americas. It is the southernmost, isthmian portion of the North American continent, which connects with South America on the southeast. When considered part of the unified continental model, it is considered a subcontinent...

 and the extreme north-western portion of South America
South America
South America is a continent situated in the Western Hemisphere, mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere. The continent is also considered a subcontinent of the Americas. It is bordered on the west by the Pacific Ocean and on the north and east...

, the white-headed capuchin is important to rainforest
Rainforest
Rainforests are forests characterized by high rainfall, with definitions based on a minimum normal annual rainfall of 1750-2000 mm...

 ecology for its role in dispersing seeds and pollen
Pollen
Pollen is a fine to coarse powder containing the microgametophytes of seed plants, which produce the male gametes . Pollen grains have a hard coat that protects the sperm cells during the process of their movement from the stamens to the pistil of flowering plants or from the male cone to the...

.

Among the best known monkey
Monkey
A monkey is a primate, either an Old World monkey or a New World monkey. There are about 260 known living species of monkey. Many are arboreal, although there are species that live primarily on the ground, such as baboons. Monkeys are generally considered to be intelligent. Unlike apes, monkeys...

s, the white-headed capuchin is recognized as the typical companion to the organ grinder.
Encyclopedia
The white-headed capuchin (Cebus capucinus), also known as the white-faced capuchin or white-throated capuchin, is a medium-sized New World monkey
New World monkey
New World monkeys are the five families of primates that are found in Central and South America: Callitrichidae, Cebidae, Aotidae, Pitheciidae, and Atelidae. The five families are ranked together as the Platyrrhini parvorder and the Ceboidea superfamily, which are essentially synonymous since...

 of the family Cebidae
Cebidae
The Cebidae is one of the five families of New World monkeys now recognised. It includes the capuchin monkeys and squirrel monkeys. These species are found throughout tropical and subtropical South and Central America.-Characteristics:...

, subfamily Cebinae. Native to the forests of Central America
Central America
Central America is the central geographic region of the Americas. It is the southernmost, isthmian portion of the North American continent, which connects with South America on the southeast. When considered part of the unified continental model, it is considered a subcontinent...

 and the extreme north-western portion of South America
South America
South America is a continent situated in the Western Hemisphere, mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere. The continent is also considered a subcontinent of the Americas. It is bordered on the west by the Pacific Ocean and on the north and east...

, the white-headed capuchin is important to rainforest
Rainforest
Rainforests are forests characterized by high rainfall, with definitions based on a minimum normal annual rainfall of 1750-2000 mm...

 ecology for its role in dispersing seeds and pollen
Pollen
Pollen is a fine to coarse powder containing the microgametophytes of seed plants, which produce the male gametes . Pollen grains have a hard coat that protects the sperm cells during the process of their movement from the stamens to the pistil of flowering plants or from the male cone to the...

.

Among the best known monkey
Monkey
A monkey is a primate, either an Old World monkey or a New World monkey. There are about 260 known living species of monkey. Many are arboreal, although there are species that live primarily on the ground, such as baboons. Monkeys are generally considered to be intelligent. Unlike apes, monkeys...

s, the white-headed capuchin is recognized as the typical companion to the organ grinder. In recent years the species has become popular in North America
North America
North America is a continent wholly within the Northern Hemisphere and almost wholly within the Western Hemisphere. It is also considered a northern subcontinent of the Americas...

n 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, invertebrate
Invertebrate
An invertebrate is an animal without a backbone. The group includes 97% of all animal species – all animals except those in the chordate subphylum Vertebrata .Invertebrates form a paraphyletic group...

s, and small vertebrate
Vertebrate
Vertebrates are animals that are members of the subphylum Vertebrata . Vertebrates are the largest group of chordates, with currently about 58,000 species described. Vertebrates include the jawless fishes, bony fishes, sharks and rays, amphibians, reptiles, mammals, and birds...

s. 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 fur 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

The white-headed capuchin was one of the many species originally described by Linnaeus
Carolus Linnaeus
Carl Linnaeus , also known after his ennoblement as , was a Swedish botanist, physician, and zoologist, who laid the foundations for the modern scheme of binomial nomenclature. He is known as the father of modern taxonomy, and is also considered one of the fathers of modern ecology...

 in his 18th century work, Systema Naturae
Systema Naturae
The book was one of the major works of the Swedish botanist, zoologist and physician Carolus Linnaeus. The first edition was published in 1735...

. It is a member of the family Cebidae
Cebidae
The Cebidae is one of the five families of New World monkeys now recognised. It includes the capuchin monkeys and squirrel monkeys. These species are found throughout tropical and subtropical South and Central America.-Characteristics:...

, the family of New World monkeys containing capuchin monkey
Capuchin monkey
The capuchins are New World monkeys of the genus Cebus. The range of capuchin monkeys includes Central America and South America as far south as northern Argentina...

s, squirrel monkey
Squirrel monkey
The squirrel monkeys are the New World monkeys of the genus Saimiri. They are the only genus in the subfamily Saimirinae.Squirrel monkeys live in the tropical forests of Central and South America in the canopy layer. Most species have parapatric or allopatric ranges in the Amazon, while S...

s, tamarin
Tamarin
The tamarins are squirrel-sized New World monkeys from the family Callitrichidae in the genus Saguinus. They are closely related to the lion tamarins in the genus Leontopithecus.- Range :...

s and marmoset
Marmoset
Marmosets are the 22 New World monkey species of the genera Callithrix, Cebuella, Callibella, and Mico. All four genera are part of the biological family Callitrichidae. The term marmoset is also used in reference to the Goeldi's Monkey, Callimico goeldii, which is closely related.Most marmosets...

s. It is the type species
Type species
In biological nomenclature, a type species is both a concept and a practical system which is used in the classification and nomenclature of animals and plants. The value of a "type species" lies in the fact that it makes clear what is meant by a particular genus name. A type species is the species...

 for the genus Cebus
CEBus
CEBus, short for Consumer Electronics Bus, also known as EIA-600, is a set of electrical standards and communication protocols for electronic devices to transmit commands and data...

, the genus that includes all the capuchin monkeys. It is a member of the C. capucinus species group within the genus Cebus, a group that also includes the white-fronted capuchin
White-fronted Capuchin
The white-fronted capuchin, Cebus albifrons, is a species of capuchin monkey, a type of New World primate, found in seven different countries in South America: Bolivia, Brazil, Colombia, Venezuela, Ecuador, Peru, and Trinidad and Tobago...

, the weeper capuchin
Weeper Capuchin
The weeper capuchin is a capuchin monkey from South America. It is found in northern Brazil, Guyana, French Guiana, Suriname, Venezuela and possibly northern Colombia.-Diet:...

 and the Kaapori capuchin
Kaapori Capuchin
The Kaapori capuchin is a capuchin monkey endemic to Brazil. This species is found in the Brazilian states of Pará and Maranhão. Formerly considered a subspecies of the weeper capuchin , it was recently elevated to species status....

.

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:
  • C. c. capucinus, from the southern part of the range in Ecuador
    Ecuador
    Ecuador , officially the Republic of Ecuador is a representative democratic republic in South America, bordered by Colombia on the north, Peru on the east and south, and by the Pacific Ocean to the west. It is one of only two countries in South America, along with Chile, that do not have a border...

    , Colombia
    Colombia
    Colombia, officially the Republic of Colombia , is a unitary constitutional republic comprising thirty-two departments. The country is located in northwestern South America, bordered to the east by Venezuela and Brazil; to the south by Ecuador and Peru; to the north by the Caribbean Sea; to the...

     and eastern Panama
    Panama
    Panama , officially the Republic of Panama , is the southernmost country of Central America. Situated on the isthmus connecting North and South America, it is bordered by Costa Rica to the northwest, Colombia to the southeast, the Caribbean Sea to the north and the Pacific Ocean to the south. The...

  • C. c. imitator, from most of Nicaragua
    Nicaragua
    Nicaragua is the largest country in the Central American American isthmus, bordered by Honduras to the north and Costa Rica to the south. The country is situated between 11 and 14 degrees north of the Equator in the Northern Hemisphere, which places it entirely within the tropics. The Pacific Ocean...

    , Costa Rica
    Costa Rica
    Costa Rica , officially the Republic of Costa Rica is a multilingual, multiethnic and multicultural country in Central America, bordered by Nicaragua to the north, Panama to the southeast, the Pacific Ocean to the west and the Caribbean Sea to the east....

     and western Panama
  • C. c. limitaneus, from Honduras
    Honduras
    Honduras is a republic in Central America. It was previously known as Spanish Honduras to differentiate it from British Honduras, which became the modern-day state of Belize...

     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.

Physical description

Like other monkeys in the genus Cebus
CEBus
CEBus, short for Consumer Electronics Bus, also known as EIA-600, is a set of electrical standards and communication protocols for electronic devices to transmit commands and data...

, the white-headed capuchin is named after the order of Capuchin friars
Order of Friars Minor Capuchin
The Order of Friars Minor Capuchin is an Order of friars in the Catholic Church, among the chief offshoots of the Franciscans. The worldwide head of the Order, called the Minister General, is currently Father Mauro Jöhri.-Origins :...

 – the cowl
Cowl
This article is about the garment used by monks and nuns. For other uses, see Cowl or Cowling .The cowl is an item of clothing consisting of a long, hooded garment with wide sleeves. Originally it may have referred simply to the hooded portion of a cloak...

s worn by these friars closely resemble the monkey's head coloration. The white-headed capuchin has mostly black fur, with white to yellowish fur on the neck, throat, chest, shoulders, and upper arms. The face is pink. A V-shaped area of black fur on the crown of the head is distinctive. It has a prehensile tail that is often held coiled, giving the white-headed capuchin the nickname "ringtail".

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). The tail is longer than the body, at up to 551 mm (21.7 in) in length. Males are about 27% larger than females. The brain
Brain
The brain is the center of the nervous system in all vertebrate and most invertebrate animals—only a few primitive invertebrates such as sponges, jellyfish, sea squirts and starfishes do not have one. It is located in the head, usually close to primary sensory apparatus such as vision, hearing,...

 of a white-headed capuchin is about 79.2 g (2.8 oz), which is larger than that of several larger monkey species, such as the Mantled Howler
Mantled Howler
The mantled howler , or golden-mantled howling monkey, is a species of howler monkey, a type of New World monkey, from Central and South America. It is one of the monkey species most often seen and heard in the wild in Central America...

.

Social structure

The white-headed capuchin is a diurnal
Diurnal animal
Diurnality is a plant or animal behavior characterized by activity during the day and sleeping at night.-In animals:Animals that are not diurnal might be nocturnal or crepuscular . Many animal species are diurnal, including many mammals, insects, reptiles and birds...

 and arboreal animal. However, it does come down to the ground more often than many other New World monkeys. It moves primarily by walking on all four limbs. It
lives in troops, or groups, of up to 40 monkeys (mean 16, range 4-40) and has a male/female adult sex ratio of .71 on average (range .54-.88). With rare exceptions, females spend their entire lives with their female kin. 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. The median age of migration in the Santa Rosa population is 4.5 years. Males sometimes migrate alone, but more often they migrate in the company of other males who are often their kin. 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. Alpha males have been known to keep their positions as long as 17 years in this species 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. 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. 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.

Kinship is an important organizing factor in the structuring of female-female social relationships. 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. 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. Female-female dyads groom far more than male-female and male-male dyads. 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.

Female capuchins have linear dominance hierarchies. 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. Males are typically dominant to females. The alpha male is always easy to discern, but there are sometimes ambiguous rankings among subordinate males. 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. Males cooperate in coalitions against potential predators, and also in defense of the group against other males. Occasionally male coalitionary aggression becomes so violent that males are killed, particularly if they are encountered roaming the forest unaccompanied by allies. 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. 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.

Immigrating males often kill young infants when they take over a group. Females band together to defend their infants from infanticidal males, but they rarely succeed in saving their infants. 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. 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. The alpha male helps defend females from subordinate males within the group as well as from infanticidal males from other groups.

Interactions between groups

White-faced capuchin troops occupy home ranges of between 32 and 86 ha (79.1 and 212.5 acre). They travel between 1 and 3 km (0.621372736649807 and 1.9 mi) daily, averaging 2 kilometres (1.2 mi) per day. 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.

Home ranges overlap extensively, 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. 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. Alpha males, who have the largest reproductive stake in the group, participate at a higher rate than subordinate males. 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.

Interspecific interactions

The white-headed capuchin sometimes interacts with other sympatric monkey species. White-headed capuchins sometimes travel with and even groom Geoffroy's Spider Monkey
Geoffroy's Spider Monkey
Geoffroy's spider monkey, Ateles geoffroyi, also known as black-handed spider monkey, is a species of spider monkey, a type of New World monkey, from Central America, parts of Mexico and possibly a small portion of Colombia. There are at least five subspecies. Some primatologists classify the...

s. However, aggressive interactions between the capuchins and spider monkeys also occur. Interactions between the white-headed capuchin and mantled howler
Mantled Howler
The mantled howler , or golden-mantled howling monkey, is a species of howler monkey, a type of New World monkey, from Central and South America. It is one of the monkey species most often seen and heard in the wild in Central America...

 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 capuchin species often travel with and feed together with squirrel monkey
Squirrel monkey
The squirrel monkeys are the New World monkeys of the genus Saimiri. They are the only genus in the subfamily Saimirinae.Squirrel monkeys live in the tropical forests of Central and South America in the canopy layer. Most species have parapatric or allopatric ranges in the Amazon, while S...

s, the white-headed capuchin only rarely associates with the Central American squirrel monkey
Central American Squirrel Monkey
The Central American squirrel monkey is a squirrel monkey species from the Pacific coast of Costa Rica and Panama. It is restricted to the northwestern tip of Panama near the border with Costa Rica, and the central and southern Pacific coast of Costa Rica, primarily in Manuel Antonio and Corcovado...

. 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.

Several non-primate animal species tend to follow troops of White-faced Monkeys or are otherwise attracted by their presence. white-lipped peccaries
White-lipped Peccary
The White-lipped Peccary, Tayassu pecari, is a peccary species found in Central and South America, living in rainforest, dry forest and chaco scrub. It is monotypic within the genus Tayassu....

 and common agouti
Common agouti
The popular term Agouti designates several rodent species of the genus Dasyprocta that inhabit areas of Middle America, the West Indies, and northern South America. They are related to guinea pigs and look quite similar but have longer legs. The species vary in color from tawny to dark brown with...

s 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
Double-toothed Kite
The Double-toothed Kite is a species of bird of prey in the Accipitridae family. It is found in Belize, Bolivia, Brazil, Colombia, Costa Rica, Ecuador, El Salvador, French Guiana, Guatemala, Guyana, Honduras, Mexico, Nicaragua, Panama, Peru, Suriname, Trinidad and Tobago, and Venezuela.This fairly...

, the white hawk
White Hawk
The White Hawk , a bird of prey breeding in the tropical New World, belongs to the family Accipitridae of the Falconiformes; it is sometimes separated in the Accipitriformes with the other hawks and their relatives...

 and the sharp-shinned hawk
Sharp-shinned Hawk
The Sharp-shinned Hawk is a small hawk. In fact, "sharp-shins" or "sharpies" are the smallest to reside in USA and Canada, though some Neotropical species are smaller...

.

Diet

The white-headed capuchin is an omnivore
Omnivore
Omnivores are species that eat both plants and animals as their primary food source...

. Its primary foods are fruit and insects. It forages at all levels of the forest, and also forages on the ground. 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 anvil
Anvil
An anvil is a basic tool, a block with a hard surface on which another object is struck. The inertia of the anvil allows the energy of the striking tool to be transferred to the work piece. In most cases the anvil is used as a forging tool...

s to crack hard fruits. Its prehensile tail assists with feeding, helping support the monkey when foraging for food below the branches.

Fruit can make up between 50% and 67% or more of the capuchin's diet. In one study in Panama
Panama
Panama , officially the Republic of Panama , is the southernmost country of Central America. Situated on the isthmus connecting North and South America, it is bordered by Costa Rica to the northwest, Colombia to the southeast, the Caribbean Sea to the north and the Pacific Ocean to the south. The...

, white-headed capuchins ate 95 different fruit species. Among its favorite fruits are figs from the family Moraceae
Moraceae
Moraceae — often called the mulberry family or fig family — are a family of flowering plants comprising about 40 genera and over 1000 species. Most are widespread in tropical and subtropical regions, less so in temperate climates...

, mangos and related fruits from the family Anacardiaceae
Anacardiaceae
Anacardiaceae are a family of flowering plants bearing fruits that are drupes and in some cases producing urushiol, an irritant. Anacardiaceae include numerous genera with several of economic importance. Notable plants in this family include cashew , mango, poison ivy, sumac, smoke tree, and marula...

, the bean-like fruits from the family Leguminosae and fruits from the family Rubiaceae
Rubiaceae
The Rubiaceae is a family of flowering plants, variously called the coffee family, madder family, or bedstraw family. The group contains many commonly known plants, including the economically important coffee , quinine , and gambier , and the horticulturally valuable madder , west indian jasmine ,...

. It generally only eats ripe fruit, testing for ripeness by smelling, tasting and prodding the fruit. It typically eats 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. It also uses the bromelids as a water source, drinking the water that gets trapped inside.

Insect prey eaten includes beetle
Beetle
Coleoptera is an order of insects commonly called beetles. The word "coleoptera" is from the Greek , koleos, "sheath"; and , pteron, "wing", thus "sheathed wing". Coleoptera contains more species than any other order, constituting almost 25% of all known life-forms...

 larvae, butterfly and moth caterpillar
Caterpillar
Caterpillars are the larval form of members of the order Lepidoptera . They are mostly herbivorous in food habit, although some species are insectivorous. Caterpillars are voracious feeders and many of them are considered to be pests in agriculture...

s, ant
Ant
Ants are social insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. More than...

s, wasp
Wasp
The term wasp is typically defined as any insect of the order Hymenoptera and suborder Apocrita that is neither a bee nor an ant. Almost every pest insect species has at least one wasp species that preys upon it or parasitizes it, making wasps critically important in natural control of their...

s, and ant and wasp larvae. It also eats larger prey, such as birds, bird eggs, frog
Frog
Frogs are amphibians in the order Anura , formerly referred to as Salientia . Most frogs are characterized by a short body, webbed digits , protruding eyes and the absence of a tail...

s, lizard
Lizard
Lizards are a widespread group of squamate reptiles, with nearly 3800 species, ranging across all continents except Antarctica as well as most oceanic island chains...

s, crab
Crab
True crabs are decapod crustaceans of the infraorder Brachyura, which typically have a very short projecting "tail" , or where the reduced abdomen is entirely hidden under the thorax...

s, mollusks and small mammals. The population in Guanacaste
Guanacaste Province
Guanacaste is a province of Costa Rica located in the northwestern part of the country, along the coast of the Pacific Ocean. To the north it borders Nicaragua. To the east is the Alajuela Province, and to the southeast is the Puntarenas Province. It is the most sparsely populated of all the...

, Costa Rica
Costa Rica
Costa Rica , officially the Republic of Costa Rica is a multilingual, multiethnic and multicultural country in Central America, bordered by Nicaragua to the north, Panama to the southeast, the Pacific Ocean to the west and the Caribbean Sea to the east....

 in particular is noted for hunting squirrel
Squirrel
Squirrels belong to a large family of small or medium-sized rodents called the Sciuridae. The family includes tree squirrels, ground squirrels, chipmunks, marmots , flying squirrels, and prairie dogs. Squirrels are indigenous to the Americas, Eurasia, and Africa and have been introduced to Australia...

s, magpie
Magpie
Magpies are passerine birds of the crow family, Corvidae.In Europe, "magpie" is often used by English speakers as a synonym for the European Magpie, as there are no other magpies in Europe outside Iberia...

s, White-crowned Parrot
White-crowned Parrot
The White-crowned Parrot , also known as the White-crowned Pionus in aviculture, is a small parrot which is a resident breeding species from eastern Mexico to western Panama....

s and baby coati
Coati
Coatis, genera Nasua and Nasuella, also known as the Brazilian aardvark, Mexican tejón, hog-nosed coon, pizotes, crackoons and snookum bears, are members of the raccoon family . They are diurnal mammals native to South America, Central America, and south-western North America...

s. The amount of vertebrate
Vertebrate
Vertebrates are animals that are members of the subphylum Vertebrata . Vertebrates are the largest group of chordates, with currently about 58,000 species described. Vertebrates include the jawless fishes, bony fishes, sharks and rays, amphibians, reptiles, mammals, and birds...

 prey eaten varies by troop. Even neighboring troops can show significant differences in their diets.

The diet can vary between the rainy and dry season. For example, in Guanacaste
Guanacaste Province
Guanacaste is a province of Costa Rica located in the northwestern part of the country, along the coast of the Pacific Ocean. To the north it borders Nicaragua. To the east is the Alajuela Province, and to the southeast is the Puntarenas Province. It is the most sparsely populated of all the...

, Costa Rica
Costa Rica
Costa Rica , officially the Republic of Costa Rica is a multilingual, multiethnic and multicultural country in Central America, bordered by Nicaragua to the north, Panama to the southeast, the Pacific Ocean to the west and the Caribbean Sea to the east....

 the white-headed capuchin can eat a wide variety of fruits as well as caterpillars in the early rainy season (June to November). But during 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. 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.

Tool use

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 monkey
Spider monkey
Spider monkeys of the genus Ateles are New World monkeys in the subfamily Atelinae, family Atelidae. Like other atelines, they are found in tropical forests of Central and South America, from southern Mexico to Brazil...

s among New World monkeys.

The white-headed capuchin is known to rub parts of certain plants into their fur. Plants used in this manner include citrus
Citrus
Citrus is a common term and genus of flowering plants in the rue family, Rutaceae. Citrus is believed to have originated in the part of Southeast Asia bordered by Northeastern India, Myanmar and the Yunnan province of China...

 fruits, vines of the genera Piper
Piper (genus)
Piper, the pepper plants or pepper vines , are an economically and ecologically important genus in the family Piperaceae...

and Clematis
Clematis
Clematis is a genus of about 300 species within the buttercup family Ranunculaceae. Their garden hybrids have been popular among gardeners beginning with Clematis × jackmanii, a garden standby since 1862; more hybrid cultivars are being produced constantly. They are mainly of Chinese and Japanese...

, monkey comb (genus Sloanea
Sloanea
Sloanea is a genus of flowering plants in the family Elaeocarpaceae, comprising about 150 species .Species include:* Sloanea acutiflora, Uittien* Sloanea assamica, Rehder & E. Wilson* Sloanea australis, Benth. & F.Muell...

), dumb cane and custard apple
Custard apple
Custard apple, a common name, can refer to:*Custard-apple, also called bull's heart, the fruit of the tree Annona reticulata.*The custard-apple tree itself, Annona reticulataThe term may also refer to:...

. Ants and millipede
Millipede
Millipedes are arthropods that have two pairs of legs per segment . Each segment that has two pairs of legs is a result of two single segments fused together as one...

s are also used in this way. It is not definitively known what this fur rubbing is for, but this may deter parasites such as tick
Tick
Ticks are small arachnids in the order Ixodida, along with mites, constitute the subclass Acarina. Ticks are ectoparasites , living by hematophagy on the blood of mammals, birds, and sometimes reptiles and amphibians...

s and insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...

s, or it may serve as a fungicide
Fungicide
Fungicides are chemical compounds or biological organisms used to kill or inhibit fungi or fungal spores. Fungi can cause serious damage in agriculture, resulting in critical losses of yield, quality and profit. Fungicides are used both in agriculture and to fight fungal infections in animals...

 or bactericide or anti-inflammatory
Anti-inflammatory
Anti-inflammatory refers to the property of a substance or treatment that reduces inflammation. Anti-inflammatory drugs make up about half of analgesics, remedying pain by reducing inflammation as opposed to opioids, which affect the central nervous system....

 agent. Alternatively, it may be a form of scent marking.

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. 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. Some populations also use trees or other hard surfaces as anvil
Anvil
An anvil is a basic tool, a block with a hard surface on which another object is struck. The inertia of the anvil allows the energy of the striking tool to be transferred to the work piece. In most cases the anvil is used as a forging tool...

s in order to crack mollusks. And it sometimes uses sticks as probes to explore openings.

The white-headed capuchin's intelligence and ability to use tools allows them to be trained to assist paraplegics. Other species of capuchin monkey
Capuchin monkey
The capuchins are New World monkeys of the genus Cebus. The range of capuchin monkeys includes Central America and South America as far south as northern Argentina...

s are also trained in this manner. White-headed capuchins can also be trained for roles on television
Television
Television is a telecommunication medium for transmitting and receiving moving images that can be monochrome or colored, with accompanying sound...

 and movies, such as Marcel on the television series Friends
Friends
Friends is an American sitcom created by David Crane and Marta Kauffman, which aired on NBC from September 22, 1994 to May 6, 2004. The series revolves around a group of friends in Manhattan. The series was produced by Bright/Kauffman/Crane Productions, in association with Warner Bros. Television...

. They were also traditionally used as organ grinder monkeys.

Communication

The white-headed capuchin is noisy. Loud calls, such as barks and coughs, are used to communicate
Animal communication
Animal communication is any behavior on the part of one animal that has an effect on the current or future behaviour of another animal. The study of animal communication, is sometimes called Zoosemiotics has played an important part in the...

 threat warnings, and softer calls, such as squeals, are used in intimate discourse. Different types of threats, such as a threat from a terrestrial animal versus a threat from a bird, invoke different vocalizations. Facial expressions and scent are also important to communication. It sometimes engages in a practice known as "urine washing", in which the monkey rubs urine on its feet. The exact purpose of this practice is unknown, but it may be a form of olfactory signal.

Reproduction

The white-headed capuchin uses a polygamous
Polygamy
Polygamy is a marriage which includes more than two partners...

 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. Although a female may mate with several males, the dominant male may be more likely to copulate when the female is at peak fertility. Nonetheless, there is evidence that dominant males do tend to avoid breeding with their own daughters who are members of the troop. Such avoidance is rare among New World primates.

Copulation takes about 2 minutes, and the gestation
Gestation
Gestation is the carrying of an embryo or fetus inside a female viviparous animal. Mammals during pregnancy can have one or more gestations at the same time ....

 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 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
Weaning
Weaning is the process of gradually introducing a mammal infant, either human or animal, to what will be its adult diet and withdrawing the supply of its mother's milk.The process takes place only in mammals, as only mammals produce milk...

 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. Capuchins engage in high levels of alloparenting
Alloparenting
In biology and sociology, alloparenting is where individuals other than the actual parents act in a parental role.One common form of alloparenting is where grandparents adopt a parental role. This is sometimes named a "skipped generation household"...

, in which monkeys other than the mother help care for the infant. Infants are carried by alloparents most often between 4 and 6 weeks in age. Males as well as females engage in alloparenting.

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

Distribution and habitat

The white-headed capuchin is found in much of Central America and a small portion of South America. In Central America
Central America
Central America is the central geographic region of the Americas. It is the southernmost, isthmian portion of the North American continent, which connects with South America on the southeast. When considered part of the unified continental model, it is considered a subcontinent...

, its range includes much of Honduras
Honduras
Honduras is a republic in Central America. It was previously known as Spanish Honduras to differentiate it from British Honduras, which became the modern-day state of Belize...

, Nicaragua
Nicaragua
Nicaragua is the largest country in the Central American American isthmus, bordered by Honduras to the north and Costa Rica to the south. The country is situated between 11 and 14 degrees north of the Equator in the Northern Hemisphere, which places it entirely within the tropics. The Pacific Ocean...

, Costa Rica
Costa Rica
Costa Rica , officially the Republic of Costa Rica is a multilingual, multiethnic and multicultural country in Central America, bordered by Nicaragua to the north, Panama to the southeast, the Pacific Ocean to the west and the Caribbean Sea to the east....

 and Panama
Panama
Panama , officially the Republic of Panama , is the southernmost country of Central America. Situated on the isthmus connecting North and South America, it is bordered by Costa Rica to the northwest, Colombia to the southeast, the Caribbean Sea to the north and the Pacific Ocean to the south. The...

. It has also been reported to occur in eastern Guatemala
Guatemala
Guatemala is a country in Central America bordered by Mexico to the north and west, the Pacific Ocean to the southwest, Belize to the northeast, the Caribbean to the east, and Honduras and El Salvador to the southeast...

 and southern Belize
Belize
Belize is a constitutional monarchy and the northernmost country in Central America. Belize has a diverse society, comprising many cultures and languages. Even though Kriol and Spanish are spoken among the population, Belize is the only country in Central America where English is the official...

, but these reports are unconfirmed. In South America the white-headed capuchin is found in the extreme north-western strip between the Pacific Ocean
Pacific Ocean
The Pacific Ocean is the largest of the Earth's oceanic divisions. It extends from the Arctic in the north to the Southern Ocean in the south, bounded by Asia and Australia in the west, and the Americas in the east.At 165.2 million square kilometres in area, this largest division of the World...

 and the Andes Mountains in Colombia
Colombia
Colombia, officially the Republic of Colombia , is a unitary constitutional republic comprising thirty-two departments. The country is located in northwestern South America, bordered to the east by Venezuela and Brazil; to the south by Ecuador and Peru; to the north by the Caribbean Sea; to the...

 and northwestern Ecuador
Ecuador
Ecuador , officially the Republic of Ecuador is a representative democratic republic in South America, bordered by Colombia on the north, Peru on the east and south, and by the Pacific Ocean to the west. It is one of only two countries in South America, along with Chile, that do not have a border...

. It is among the most commonly seen monkeys in Central America's national parks, such as Manuel Antonio National Park
Manuel Antonio National Park
Manuel Antonio National Park, in Spanish the Parque Nacional Manuel Antonio, is a small National Park in the Central Pacific Conservation Area located on the Pacific coast of Costa Rica, just south of the city of Quepos, Puntarenas, and from the national capital of San José...

, Corcovado National Park
Corcovado National Park
Corcovado National Park is a National Park on the Osa Peninsula in the South West of Costa Rica , which is part of the Osa Conservation Area. It was established on 24 October 1975, and encompasses an area of 425 km² . It is widely considered the crown jewel in the extensive system of national...

, Santa Rosa National Park
Santa Rosa National Park
Santa Rosa National Park, in Spanish the , is a national park, part of the Area de Conservación Guanacaste World Heritage site, in the northern western part of Costa Rica, originally created to protect the scene of the Battle of Santa Rosa, at its eastern edge lies the Interamerican Highway where...

 and Soberania National Park
Soberanía National Park
Soberanía National Park is a national park in Panama near the banks of the Panama Canal in the provinces of Panamá and Colón, some from Panama City. The Chagres River runs through the park....

.

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
Mangrove
Mangroves are various kinds of trees up to medium height and shrubs that grow in saline coastal sediment habitats in the tropics and subtropics – mainly between latitudes N and S...

 and montane
Montane
In biogeography, montane is the highland area located below the subalpine zone. Montane regions generally have cooler temperatures and often have higher rainfall than the adjacent lowland regions, and are frequently home to distinct communities of plants and animals.The term "montane" means "of the...

 forests. However, it appears to prefer primary or advanced secondary forest
Secondary forest
A secondary forest is a forest or woodland area which has re-grown after a major disturbance such as fire, insect infestation, timber harvest or windthrow, until a long enough period has passed so that the effects of the disturbance are no longer evident...

s. 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.

Conservation status

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. Its status can also be harmed by deforestation
Deforestation
Deforestation is the removal of a forest or stand of trees where the land is thereafter converted to a nonforest use. Examples of deforestation include conversion of forestland to farms, ranches, or urban use....

. 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. 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. The white-headed capuchin is important to its ecosystems as a seed and pollen
Pollen
Pollen is a fine to coarse powder containing the microgametophytes of seed plants, which produce the male gametes . Pollen grains have a hard coat that protects the sperm cells during the process of their movement from the stamens to the pistil of flowering plants or from the male cone to the...

 disperser. It also impacts the ecosystem by eating insects that act as pests to certain trees, by pruning certain trees, such as Gustavia superba
Gustavia superba
Gustavia superba is a species of trees that grow in Central and North-Western America.Common names include Membrillo, Sachamango and Heaven Lotus. These plants are noted for having very long leaves and large flowers . The trunks themselves are mostly 6-10m high with the leaves radiating from the top...

and Bursera simaruba
Bursera simaruba
Bursera simaruba, commonly known as the Gumbo-limbo, is a tree species in the family Burseraceae, native to tropical regions of the Americas from the southeasternmost United States south through Mexico and the Caribbean to Brazil and Venezuela...

, causing them generate more branches and possibly additional fruit, and by accelerating germination
Germination
Germination is the process in which a plant or fungus emerges from a seed or spore, respectively, and begins growth. The most common example of germination is the sprouting of a seedling from a seed of an angiosperm or gymnosperm. However the growth of a sporeling from a spore, for example the...

 of certain seeds when they pass through the capuchin's digestive tract. In addition, the white-headed capuchin sometimes kills Acacia collinsii
Acacia collinsii
Acacia collinsii is a species of flowering plant. It grows in secondary succession in seasonally dry ecosystems in southern Central America, where it exhibits a symbiotic relationship with several species of ant...

plants when it rips through the plant's branches to get to resident ant colonies.

External links

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