White-fronted Capuchin
Encyclopedia
The white-fronted capuchin, Cebus albifrons, is a 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...

, a type of New World
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...

 primate
Primate
A primate is a mammal of the order Primates , which contains prosimians and simians. Primates arose from ancestors that lived in the trees of tropical forests; many primate characteristics represent adaptations to life in this challenging three-dimensional environment...

, found in seven different countries in 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...

: Bolivia
Bolivia
Bolivia officially known as Plurinational State of Bolivia , is a landlocked country in central South America. It is the poorest country in South America...

, Brazil
Brazil
Brazil , officially the Federative Republic of Brazil , is the largest country in South America. It is the world's fifth largest country, both by geographical area and by population with over 192 million people...

, 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...

, Venezuela
Venezuela
Venezuela , officially called the Bolivarian Republic of Venezuela , is a tropical country on the northern coast of South America. It borders Colombia to the west, Guyana to the east, and Brazil to the south...

, 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...

, Peru
Peru
Peru , officially the Republic of Peru , is a country in western South America. It is bordered on the north by Ecuador and Colombia, on the east by Brazil, on the southeast by Bolivia, on the south by Chile, and on the west by the Pacific Ocean....

, and Trinidad and Tobago
Trinidad and Tobago
Trinidad and Tobago officially the Republic of Trinidad and Tobago is an archipelagic state in the southern Caribbean, lying just off the coast of northeastern Venezuela and south of Grenada in the Lesser Antilles...

. The species is divided into several different subspecies
Subspecies
Subspecies in biological classification, is either a taxonomic rank subordinate to species, ora taxonomic unit in that rank . A subspecies cannot be recognized in isolation: a species will either be recognized as having no subspecies at all or two or more, never just one...

, though the specific divisions are uncertain and controversial.

This primate is a medium-sized monkey with a light brown back and a creamy white underside. Like other 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, it is omnivorous
Omnivore
Omnivores are species that eat both plants and animals as their primary food source...

, feeding primarily on fruit
Fruit
In broad terms, a fruit is a structure of a plant that contains its seeds.The term has different meanings dependent on context. In non-technical usage, such as food preparation, fruit normally means the fleshy seed-associated structures of certain plants that are sweet and edible in the raw state,...

s, 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, other plant parts and sometimes small vertebrates. It is predated upon primarily by raptors and probably small cats, especially the Margay
Margay
The Margay is a spotted cat native to Middle and South America. Named for Prince Maximilian of Wied-Neuwied, it is a solitary and nocturnal animal that prefers remote sections of the rainforest. Although it was once believed to be vulnerable to extinction, the IUCN now lists it as "Near Threatened"...

, though snakes have been known to attack the species. It is a polygamous animal and lives on fairly large groups of 15 to 35 individuals, reproductive females give birth to a single young at biennial intervals. The species maintains a home range
Home range
Home range is the area where an animal lives and travels in. It is closely related to, but not identical with, the concept of "territory".The concept that can be traced back to a publication in 1943 by W. H. Burt, who constructed maps delineating the spatial extent or outside boundary of an...

 of 1.2 to 1.5 km² (0.463322590311042 to 0.579153237888803 sqmi) and has a complex vocal repertoire. It is one of the few primates to have been observed crafting and utilising tools in the wild.

White-fronted capuchin populations are declining. The decline is believed to be caused by human-induced habitat loss
Habitat destruction
Habitat destruction is the process in which natural habitat is rendered functionally unable to support the species present. In this process, the organisms that previously used the site are displaced or destroyed, reducing biodiversity. Habitat destruction by human activity mainly for the purpose of...

 and degradation, and hunting. In 2008, the International Union for Conservation of Nature (IUCN) classified the species as Least Concern, though subspecies in northern Colombia are considered Vulnerable. The total population of the subspecies found in Trinidad was 61 at the last census and is Critically Endangered.

Taxonomy

This species has had problems with its name, description and type locality. The holotype
Holotype
A holotype is a single physical example of an organism, known to have been used when the species was formally described. It is either the single such physical example or one of several such, but explicitly designated as the holotype...

 does not exist; the original description by Alexander von Humboldt
Alexander von Humboldt
Friedrich Wilhelm Heinrich Alexander Freiherr von Humboldt was a German naturalist and explorer, and the younger brother of the Prussian minister, philosopher and linguist Wilhelm von Humboldt...

 in 1812 describes an animal that is much darker (grayish) than those that exist close to the type locality, and the description includes a dark tail tip, a character that is completely unknown in any population of the species. Additionally, the animal which von Humboldt examined was a tame animal in Maipures, where the species is not usually found. The closest population is about three kilometers to the north, on the other side of the Tuparro River
Tuparro River
Tuparro River is a river of Colombia found in Vichada Department. It gives its name to El Tuparro National Natural Park. Tuparro is part of the Orinoco River basin....

.

Defler and Hernández established a neotype from the population that was called Cebus albifrons albifrons by Hernández C. and Cooper. Another problem has been that the taxon
Taxon
|thumb|270px|[[African elephants]] form a widely-accepted taxon, the [[genus]] LoxodontaA taxon is a group of organisms, which a taxonomist adjudges to be a unit. Usually a taxon is given a name and a rank, although neither is a requirement...

 C. a. unicolor described by Spix (1823) and further defined by Hershkovitz was indistinguishable from C. a. albifrons; the two are synonymous.

Taxonomic classification

Hershkovitz (1949) originally named 13 subspecies, while Hernández-Camacho and Cooper (1976) described eight subspecies for Colombia. Colin Groves
Colin Groves
Colin Peter Groves is Professor of Biological Anthropology at the Australian National University in Canberra, Australia.Born in England on 24 June 1942, Colin Groves completed a Bachelor of Science at University College London in 1963, and a Doctor of Philosophy at the Royal Free Hospital School of...

 assessed the species in 2001, further reducing the number. One notable subspecies outside of Colombia is the critically endangered Trinidad white-fronted capuchin. The following subspecies are recognised by assessors working for the IUCN.
  • Ecuadorian white-fronted capuchin, Cebus albifrons aequatorialis, found in Ecuador and Peru.
  • White-fronted capuchin, Cebus albifrons albifrons, found in Bolivia, Brazil, Colombia, Peru and Venezuela.
  • Río Cesar white-fronted capuchin, Cebus albifrons cesarae, found in Colombia.
  • Shock-headed capuchin, Cebus albifrons cuscinus, found in Bolivia, Brazil, Ecuador and Peru.
  • Santa Marta white-fronted capuchin, Cebus albifrons malitiosus, found along the northern flanks of the Sierra Nevada de Santa Marta
    Sierra Nevada de Santa Marta
    The Sierra Nevada de Santa Marta is an isolated mountain range apart from the Andes chain that runs through Colombia. Reaching an altitude of 5,700 metres above sea level just 42 km from the Caribbean coast, the Sierra Nevada is the world's highest coastal range...

     in Colombia.
  • Trinidad white-fronted capuchin, Cebus albifrons trinitatis, 61 individuals estimated on the island of Trinidad
    Trinidad
    Trinidad is the larger and more populous of the two major islands and numerous landforms which make up the island nation of Trinidad and Tobago. It is the southernmost island in the Caribbean and lies just off the northeastern coast of Venezuela. With an area of it is also the fifth largest in...

     in 2008.
  • Varied white-fronted capuchin, Cebus albifrons versicolor, found in Colombia.


The difficulties in identifying separate subspecies, or even species, is pronounced. Hernández-Camacho and Cooper reported some specimens of capuchin from the Barranquilla
Barranquilla
Barranquilla is an industrial port city and municipality located in northern Colombia, near the Caribbean Sea. The capital of the Atlántico Department, it is the largest industrial city and port in the Colombian Caribbean region with a population of 1,148,506 as of 2005, which makes it Colombia's...

 animal market had supposedly come from the middle valley of the San Jorge River. It is difficult to determine whether these are white-headed capuchin
White-headed Capuchin
The white-headed capuchin , also known as the white-faced capuchin or white-throated capuchin, is a medium-sized New World monkey of the family Cebidae, subfamily Cebinae...

s (Cebus capucinus) or white-fronted capuchins. Intermediate characteristics include a dark crown that is high and removed from the forehead. The white parts on the face are more distinctively bald and the outside parts of the arms and legs are more clear; this suggests they are white-headed capuchin. Some specimens of C. a. versicolor seen in the market at Magangué
Magangue
Magangué Magangué is a municipality in the Department of Bolivar in Colombia, located on the banks of the Magdalena River in the North Coast. It is the capital of the southern Department, and is the second most important town....

‚ and probably captured in the lower Cauca River
Cauca River
The Cauca River is a river in Colombia that lies between the Occidental and Central cordilleras. Born in southwestern Colombia near the city of Popayán, it joins the Magdalena River near Pinillos in Bolívar Department, and the combined river eventually flows out into the Caribbean Sea. It has a...

, show similar tendencies to the above, except that there is no increase in the dark pigmentation. Based on these observations and on various "intermediate" specimens from northern Colombia, it seems possible that an investigation of the contact zone between the white-headed capuchin and white-fronted capuchin ultimately could show that these forms are conspecific. Another critical zone for this analysis is an area in northeast Ecuador where C. a. aecuatoriales and White-headed Capuchin are found, although neither sympatric distributions or intergradation
Intergradation
In zoology, intergradation is the way in which two distinct subspecies are connected via areas where populations are found that have the characteristics of both...

 have as yet been determined.

Physical description

Male white-fronted capuchins usually weigh an average of 3.4 kg (7.5 lb) and the females an average of 2.9 kg (6.4 lb), although a male on Mirití-Paraná
Mirití-Paraná
Mirití-Paraná is a town and municipality in the southern Colombian Department of Amazonas.-References:*...

 in Colombia weighed 5.5 kg (12.1 lb). This primate is usually maroon-white or palomino
Palomino
Palomino is a coat color in horses, consisting of a gold coat and white mane and tail. Genetically, the palomino color is created by a single allele of a dilution gene called the cream gene working on a "red" base coat...

 and creamy white. It has short fingers and an opposable thumb. Like other capuchin species its premolars are large, and it has square-shaped molar with a thick enamel to help with cracking nuts. Below are descriptions of the known subspecies for 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...

.
  • Cebus albifrons albifrons is found in eastern Vichada, close to the type locality, and was defined by von Humboldt using a tame animal maintained by humans (and a pig) in the village of Maipures. The original description of von Humboldt described an ashy gray animal with a black tail tip, characteristics that are not typical of any known population of C. albifrons. The C. albifrons located three kilometers to the north of Maipures are very light colored animals with yellowish or reddish tones, very similar to the population of Arauca. Cebus a. unicolor is also very light colored with yellowish tones and it seems clear that it is a synonym of C. a. albifrons. One population of very pallid coloration is found in Arauca, the northern part of Boyacá and the eastern part of Norte de Santander
    Department of North Santander
    North Santander is a department of Colombia. It is in the north of the country, bordering Venezuela. Its capital is Cúcuta, one of the major cities of Colombia....

     and probably represents C. a. albifrons.

  • Cebus a. cesarae is very light in color and quite well-defined as a subspecies. "The cap is cinnamon or snuff-brown; median dorsal region, forearm and forelag with orangeous and contrasted with sides of back and trunk; hairs of belly and chest ochracous-orange to pale ochraceous-buff and silvery; contrasting pale area of front extending over variable amounts of upper surface of shoulder and inner side of upper arm" (Hershkovitz, 1949).

  • Cebus a. malitiosus is characterized by a color that is rather dark brown over almost the entire body with yellowish shoulders. "Pale area of front less extensive, upperparts and limbs paler than in hypoleucus. Cap prout's brown, median dorsal region cinnamon brown, forearm and foreleg not markedly contrasting in color with back and sides of body; hairs of belly and chest ochraceous-tawny to cinnamon-brown and silvery; contrasting pale area of front extending well over upper surface of shoulder and inner side of upper arm" (Hershkovitz, 1949).

  • Cebus a. versicolor is a complex which includes dark populations and lighter populations. It is thought to include the subspecies C. a. leucocephalus and C. a. pleei. Herskovitz's description of C. a. pleei is of a very reddish animal, particularly in its limbs while C. a. versicolor is a lighter red. C. a. leucocephalus was described as a dark brown animal with reddish tonalities in the hind legs. Nevertheless, Hernández-Camacho and Cooper discuss evidence that the three subspecies (C. a. leucocephalus, C. a. pleei and C. a. versicolor) could be subsumed into one subspecies (C. a. versicolor), since the variations seem to be found in a very well-defined zone and even in the same groups, close to Barrancabermeja
    Barrancabermeja
    Barrancabermeja is a city and municipality in Santander Department, in northeastern Colombia. It is located on the banks of the Magdalena River in the Middle Magdalena region, 110 km west of Bucaramanga. Founded in 1536, Barrancabermeja is home of the biggest petroleum refinery in Colombia,...

     on the eastern bank of the middle Magdalena River
    Magdalena River
    The Magdalena River is the principal river of Colombia, flowing northward about through the western half of the country. It takes its name from the biblical figure Mary Magdalene. It is navigable through much of its lower reaches, in spite of the shifting sand bars at the mouth of its delta, as...

     in the Department of Santander. This strongly suggests that the dark phase (C. a. leucocephalus) and the light phase (C. a. pleei) are extremes of an intermediate (C. a. versicolor).

  • Cebus a. cuscinus is subspecies found south of the Guamués River and colored a light brown.

Geographic range and habitat

The white-fronted capuchin is found in a variety of forest types. In Vichada it exploits a more xeric habitat in terms of drainage, compared with the tufted capuchin
Tufted Capuchin
The tufted capuchin , also known as brown capuchin or black-capped capuchin is a New World primate from South America...

, which tends to be found in forests that are more mesophytic
Mesophyte
Mesophytes are terrestrial plants which are adapted to neither a particularly dry nor particularly wet environment. An example of a mesophytic habitat would be a rural temperate meadow, which might contain Goldenrod, Clover, Oxeye Daisy, and Rosa multiflora.Mesophytes make up the largest ecological...

. It is also found in flooded forests. The White-fronted Capuchin survives well in forests growing over white sand and in forests of "high caatinga
Caatinga
Caatinga is a type of vegetation, and an ecoregion characterized by this vegetation in the northeastern part of Brazil. The name "Caatinga" is a Tupi word meaning "white forest" or "white vegetation"...

" growing in the rocks and gravel at the foot of mesa
Mesa
A mesa or table mountain is an elevated area of land with a flat top and sides that are usually steep cliffs. It takes its name from its characteristic table-top shape....

s.

In Colombia, the white-fronted capuchin is found from the northern slopes of the Sierra de Santa Marta
Sierra Nevada de Santa Marta
The Sierra Nevada de Santa Marta is an isolated mountain range apart from the Andes chain that runs through Colombia. Reaching an altitude of 5,700 metres above sea level just 42 km from the Caribbean coast, the Sierra Nevada is the world's highest coastal range...

 to the south, in the valley of the Magdalena River
Magdalena River
The Magdalena River is the principal river of Colombia, flowing northward about through the western half of the country. It takes its name from the biblical figure Mary Magdalene. It is navigable through much of its lower reaches, in spite of the shifting sand bars at the mouth of its delta, as...

 to an as yet undefined point in the Department of Tolima
Department of Tolima
Tolima is one of the 32 departments of Colombia, located in the Andean region, in the center-west of the country. It is bordered on the north and the west by the department of Caldas; on the east by the department of Cundinamarca; on the south by the department of Huila, and on the west by the...

 and in the valley of the lower Cauca River
Cauca River
The Cauca River is a river in Colombia that lies between the Occidental and Central cordilleras. Born in southwestern Colombia near the city of Popayán, it joins the Magdalena River near Pinillos in Bolívar Department, and the combined river eventually flows out into the Caribbean Sea. It has a...

, to the eastern parts of central Antioquia and the southern parts of Sucre to the west. In Guajira
Department of La Guajira
La Guajira is a department of Colombia. It occupies most of its namesake peninsula, the Guajira Peninsula in the northeast region of the country, facing the Caribbean Sea and Venezuela in the northern most part of South America...

 the species is found to Riohacha
Riohacha
Riohacha, Rio Hacha or Rio de la Hacha , is a city in the Riohacha Municipality in the northern Caribbean Region of Colombia by the mouth of the Ranchería River and the Caribbean sea, capital city of the La Guajira Department. Founded by conquistador Nikolaus Federmann in 1535, Riohacha was named...

, and an isolated population is apparently found in the Serranía de Macuira
Serranía de Macuira
Serranía de Macuira is a mountain range in northern Colombia located in the municipality of Uribia, Guajira Peninsula and part of the La Guajira Department. The Serrania de Macuira stands in the middle of the La Guajira Desert at 864 meters over sea level, isolated from the mountain ranges of...

, though this needs confirmation. The species is also found along the slopes of the Serranía de Perijá and the Cordillera Oriental. To the east of the Cordillera the species is found in Norte de Santander
Department of North Santander
North Santander is a department of Colombia. It is in the north of the country, bordering Venezuela. Its capital is Cúcuta, one of the major cities of Colombia....

, western Arauca, in eastern Vichada between the Meta
Meta River
The Meta River is formed in the Meta Department, Colombia by the confluence of the Humea, Guatiquía and Guayuriba rivers. It flows east-northeastward across the Llanos Orientales plains of Colombia through an ancient fault...

 and Tuparro rivers, and then south of the Vichada River
Vichada River
The Vichada River is a blackwater river in the country of Colombia, South America. It flows into the Orinoco River.The westward course of the Vichada is offset by an impact structure, called the Vichada Structure. The structure is most likely the largest impact structure in South America....

; although east of the Ariari River, not including the Ariari itself. It is not known whether the species is found in the rather extensive forests of the upper Manacasías River in Meta. South of the Guayabero and Guaviare River
Guaviare River
The Guaviare is a tributary of the Orinoco located in Colombia.The Guaviare has its source in two other rivers, the Ariari and the Guayabero, which in turn have their own sources in the eastern part of the Andes. At long, it is the longest river on the Orinoco and is navigable for of its total...

, the white-fronted capuchin is found throughout the Amazon
Amazon Basin
The Amazon Basin is the part of South America drained by the Amazon River and its tributaries that drains an area of about , or roughly 40 percent of South America. The basin is located in the countries of Bolivia, Brazil, Colombia, Ecuador, Guyana, Peru, and Venezuela...

. The species is known to an altitude of 1500 – in the Department of Tolima
Department of Tolima
Tolima is one of the 32 departments of Colombia, located in the Andean region, in the center-west of the country. It is bordered on the north and the west by the department of Caldas; on the east by the department of Cundinamarca; on the south by the department of Huila, and on the west by the...

.

Outside of Colombia, the white-fronted capuchin is found from the Andes
Andes
The Andes is the world's longest continental mountain range. It is a continual range of highlands along the western coast of South America. This range is about long, about to wide , and of an average height of about .Along its length, the Andes is split into several ranges, which are separated...

 throughout eastern Ecuador, Peru and northern Bolivia to the Tapajós
Tapajós
The Tapajós, a Brazilian river running through a humid and hot valley, pours into the Amazon River 500 miles above Pará and is about 1200 miles long.It rises on the lofty Brazilian plateau near Diamantino in 14 degrees 25' south latitude...

 river in Brazil
Brazil
Brazil , officially the Federative Republic of Brazil , is the largest country in South America. It is the world's fifth largest country, both by geographical area and by population with over 192 million people...

, south of the Amazon River
Amazon River
The Amazon of South America is the second longest river in the world and by far the largest by waterflow with an average discharge greater than the next seven largest rivers combined...

. North of the Amazon River the species is found in the southern parts of the Venezuelan Federal State of Amazonas
Amazonas (Venezuelan state)
Amazonas State is one of the 23 states into which Venezuela is divided.The state capital is Puerto Ayacucho. The capital until the early 1900s was San Fernando de Atabapo. Although named after the Amazon River, most of the state is drained by the Orinoco. Amazonas State covers a total surface...

 and in northern Brazil between Colombia and the Branco River
Branco River
The Rio Branco is the principal affluent of the Rio Negro from the north; it is enriched by many streams from the sierras which separate Venezuela and Guyana from Brazil. Its two upper main tributaries are the Urariquira and the Takutu...

.

Cebus albifrons albifrons is very common in the eastern half of El Tuparro National Park, Colombia. It is less common in Amacayacu National Park
Amacayacu National Park
Amacayacu National Natural Park is a national park located along the Amazon River in the Amazonas Department in the south of Colombia. The park was created in 1975. The word "Amacayacu" means "River of the Hamocs" in the indigenous language Quechua. The Ticuna people currently inhabit a part of...

. Cebus a. yuracus is known south of the Putumayo River
Putumayo River
The Içá or Putumayo River is one of the tributaries of the Amazon River, west of and parallel to the Yapura. It forms part of Colombia's border with Ecuador, as well as most of the frontier with Peru...

. Cebus a. versicolor is widespread on the middle-Magdalena River
Magdalena River
The Magdalena River is the principal river of Colombia, flowing northward about through the western half of the country. It takes its name from the biblical figure Mary Magdalene. It is navigable through much of its lower reaches, in spite of the shifting sand bars at the mouth of its delta, as...

 and is observable in preserved woodlots of protected finca
Finca
Finca is a board game published by Hans im Glück. Players collect fruit by stepping around a windmill and then selling the fruit in specific combinations to the various towns on the game board.The game was nominated for the Spiel des Jahres in 2009....

s. Cebus a. malitiosus is easy to observe in Tayrona National Park, east of Santa Marta
Santa Marta
Santa Marta is the capital city of the Colombian department of Magdalena in the Caribbean Region. It was founded in July 29, 1525 by the Spanish conqueror Rodrigo de Bastidas, which makes it the oldest remaining city in Colombia...

. Cebus a. cesarae can be located in the Serranía del Perijá
Serrania del Perija
The Serranía del Perijá, Cordillera de Perijá or Sierra de Perijá is a mountain range, an extension of the eastern Andean branch , in northern South America, between Colombia and Venezuela, ending further north in the Guajira Desert, a total distance of about 310 km...

 east of Valledupar
Valledupar
Valledupar is a city and municipality in northeastern Colombia. It is the capital of Cesar Department and was founded in 1550 by the Spanish conquistador Hernando de Santana. Its name, Valle de Upar , was established in honor of the Amerindian cacique who ruled the valley; Cacique Upar...

, Cesar also in Colombia.

Behavior

This species has been studied 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...

 by Defler
Thomas Defler
Thomas R. Defler is a North American primatologist who has lived and worked in Colombia for many years. He earned his PhD from the University of Colorado at Denver in 1976 and then he moved to Colombia...

, in two different sites in Peru
Peru
Peru , officially the Republic of Peru , is a country in western South America. It is bordered on the north by Ecuador and Colombia, on the east by Brazil, on the southeast by Bolivia, on the south by Chile, and on the west by the Pacific Ocean....

 by Soini and Terborgh, in Trinidad
Trinidad
Trinidad is the larger and more populous of the two major islands and numerous landforms which make up the island nation of Trinidad and Tobago. It is the southernmost island in the Caribbean and lies just off the northeastern coast of Venezuela. With an area of it is also the fifth largest in...

 by Phillips and 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...

 by Matthews.

In eastern Vichada, Colombia, the white-fronted capuchin is found in large groups of around 35 individuals, while to the south in closed forest (perhaps as a result of competition with the tufted capuchin
Tufted Capuchin
The tufted capuchin , also known as brown capuchin or black-capped capuchin is a New World primate from South America...

) is has an average group size of 8–15 individuals. A group in Vichada used a home range of about 1.2 km² (0.463322590311042 sq mi), while Terborgh found a home range of more than 1.5 km² (0.579153237888803 sq mi) and Matthews calculated 240 hectares (593.1 acre). Near the type locality in gallery forest
Gallery forest
Gallery forests are evergreen forests that form as corridors along rivers or wetlands and project into landscapes that are otherwise only sparsely treed such as savannas, grasslands or deserts....

 and islands of forest in Vichada, the species has an ecological density of around 30 individuals/km². In forests with closed-canopy in Colombia and in southern Vichada, many areas have very low densities. Around the lower Apaporis River, for example, densities are less than one individual/km² and the size of the groups is around 15 individuals. Low densities in many parts of the Colombian Amazon make it difficult to detect the presence of the species in many parts.

Terborgh found an average of 1800 metres (1.1 mi) for the day range of a group, and calculated the following time budget of the study group in Manú National Park
Manú National Park
Manú National Park is a biosphere reserve located in Madre de Dios and Paucartambo, Cusco. Before becoming an area protected by the Peruvian government, the Manú National Park was conserved thanks to its inaccessibility. The park remains fairly inaccessible by road to this day. In 1977, UNESCO...

, Peru: 18% rest, 21% travel, 22% feeding on plant material and 39% feeding on insects; total feeding 61%. Matthews however, registered 54% foraging, 25% moving and 21% feeding and socializing. It is primarily quadrupedal, although it utilizes a great variety of gallops, jumps, falls and climbing. During certain times of the year the white-fronted capuchin is extremely terrestrial, especially when there is a scarcity of available fruits and the troop must search for arthropod
Arthropod
An arthropod is an invertebrate animal having an exoskeleton , a segmented body, and jointed appendages. Arthropods are members of the phylum Arthropoda , and include the insects, arachnids, crustaceans, and others...

s in the dry leaves of the forest floor. In some parts of the Llanos Orientales
Llanos
The Llanos is a vast tropical grassland plain situated to the east of the Andes in Colombia and Venezuela, in northwestern South America. It is an ecoregion of the Flooded grasslands and savannas Biome....

 it is found walking over the grassy savanna
Savanna
A savanna, or savannah, is a grassland ecosystem characterized by the trees being sufficiently small or widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach the ground to support an unbroken herbaceous layer consisting primarily of C4 grasses.Some...

 between forests, leaving well-beaten trails. In Vichada it uses preferential trees for sleeping at heights of 25 –. The palm Attalea
Attalea (genus)
Attalea is a large genus of palms native to Mexico, the Caribbean, Central and South America. This pinnately leaved, non-spiny genus includes both small palms lacking an aboveground stem and large trees. The genus has a complicated taxonomic history, and has often been split into four or five...

 regia
is often used for sleeping in this zone.

Diet

All species of capuchin tend to have a rather similar diet in broad terms; they are omnivores, eating fruits and small invertebrates, small vertebrates and birds' eggs, which they forage at all levels of the forest, frequently descending to the forest floor. In northern Colombia during the dry season when there are few fruits to be found, the white-fronted capuchin spends more than half its time on the ground, searching for and capturing small prey
Predation
In ecology, predation describes a biological interaction where a predator feeds on its prey . Predators may or may not kill their prey prior to feeding on them, but the act of predation always results in the death of its prey and the eventual absorption of the prey's tissue through consumption...

. It is extremely good at manipulating objects, and spends a great deal of time examining dry leaves from which it collects invertebrates (for example small 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...

s and 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' eggs) from rolled up leaves. It hunts 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 and drinks the water which accumulates in the spaces between the bracteoles of the common plant Phenakospermum
Phenakospermum
Phenakospermum is a monotypic genus in the family Strelitziaceae. Only one species is recognized, Phenakospermum guianense, native to Surinam and the eastern Amazon River basin...

 guianense
, where the frogs hide. Hunting amphibian
Amphibian
Amphibians , are a class of vertebrate animals including animals such as toads, frogs, caecilians, and salamanders. They are characterized as non-amniote ectothermic tetrapods...

s seems to be a cultural phenomenon which the members of each group learn. P. guianense is commonly present in large, dense stands in some types of forest.

In Manú National Park the animal material in the diet includes frogs, 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, small mammals and birds' eggs as well as many invertebrates, including orthoptera
Orthoptera
Orthoptera is an order of insects with paurometabolous or incomplete metamorphosis, including the grasshoppers, crickets and locusts.Many insects in this order produce sound by rubbing their wings against each other or their legs, the wings or legs containing rows of corrugated bumps...

ns, lepidoptera
Lepidoptera
Lepidoptera is a large order of insects that includes moths and butterflies . It is one of the most widespread and widely recognizable insect orders in the world, encompassing moths and the three superfamilies of butterflies, skipper butterflies, and moth-butterflies...

ns and hymenoptera
Hymenoptera
Hymenoptera is one of the largest orders of insects, comprising the sawflies, wasps, bees and ants. There are over 130,000 recognized species, with many more remaining to be described. The name refers to the heavy wings of the insects, and is derived from the Ancient Greek ὑμήν : membrane and...

ns (especially ants and 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...

 larva
Larva
A larva is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle...

e). In the Pacaya–Samiria National Reservation, it has been observed eating tent caterpillar
Tent caterpillar
Tent caterpillars are moderately sized species in the genus Malacosoma and in the moth family Lasiocampidae. Species occur in North America, Mexico, and Eurasia. Twenty-six species have been described, six of which occur in North America. Some species are considered to have subspecies as well. ...

s. Terborgh identified 73 species of plants from 33 families consumed by this primate. The 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...

 was the most important family by a wide margin, counting the number of species (17) eaten, equivalent to 23.3% of all plant species consumed. Importance values for plant families consumed by the White-fronted Capuchin in one study are as follows: Moraceae (17, 23.3%); Leguminosae
Fabaceae
The Fabaceae or Leguminosae, commonly known as the legume, pea, or bean family, is a large and economically important family of flowering plants. The group is the third largest land plant family, behind only the Orchidaceae and Asteraceae, with 730 genera and over 19,400 species...

 (5, 6.8%); Araceae
Araceae
Araceae are a family of monocotyledonous flowering plants in which flowers are borne on a type of inflorescence called a spadix. The spadix is usually accompanied by, and sometimes partially enclosed in, a spathe or leaf-like bract. Also known as the Arum family, members are often colloquially...

 (4, 5.5%); Bombacaceae
Bombacaceae
Bombacaceae is a family of flowering plants or Angiospermae included within Malvales order. As is true for any botanical name, circumscription and status of the taxon has varied with taxonomic point of view...

 (4, 5.5%); Palmae
Arecaceae
Arecaceae or Palmae , are a family of flowering plants, the only family in the monocot order Arecales. There are roughly 202 currently known genera with around 2600 species, most of which are restricted to tropical, subtropical, and warm temperate climates...

 (4, 5.5%).

Defler collected 40 species of plants from 23 families eaten by this primate in Vichada according to species consumed per family: Arecaceae
Arecaceae
Arecaceae or Palmae , are a family of flowering plants, the only family in the monocot order Arecales. There are roughly 202 currently known genera with around 2600 species, most of which are restricted to tropical, subtropical, and warm temperate climates...

 (7); 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...

 (6); Chrysobalanaceae
Chrysobalanaceae
Chrysobalanaceae is a family of trees, shrubs and flowering plants, consisting of 17 genera and about 460 species of leptocaul that grows in the Tropics or is subtropical and common in the Americas...

 (3); Leguminosae
Fabaceae
The Fabaceae or Leguminosae, commonly known as the legume, pea, or bean family, is a large and economically important family of flowering plants. The group is the third largest land plant family, behind only the Orchidaceae and Asteraceae, with 730 genera and over 19,400 species...

 (3); Passifloraceae
Passifloraceae
Passifloraceae is a family of flowering plants, containing about 530 species classified in around 18 genera. They include trees, shrubs, lianas and climbing plants, and are mostly found in tropical regions....

 (2); Bromeliaceae
Bromeliaceae
Bromeliaceae is a family of monocot flowering plants of around 3,170 species native mainly to the tropical Americas, with a few species found in the American subtropics and one in tropical west Africa, Pitcairnia feliciana...

 (2); Burseraceae
Burseraceae
Burseraceae is a moderate-sized family of 17-18 genera and about 540 species of flowering plants. The actual numbers differ according to the time period in which a given source is written describing this family. The Burseraceae is also known as the Torchwood family, the frankincense and myrrh...

 (2); Bombacaceae
Bombacaceae
Bombacaceae is a family of flowering plants or Angiospermae included within Malvales order. As is true for any botanical name, circumscription and status of the taxon has varied with taxonomic point of view...

 (1); Celastraceae
Celastraceae
The Celastraceae , is a family of about 90-100 genera and 1,300 species of vines, shrubs and small trees, belonging to the order Celastrales...

 (1); Connaraceae
Connaraceae
The Connaraceae is a family of 16 genera and about 350 species.-Genera:* Agelaea* Burttia* Cnestidium* Cnestis* Connarus* Ellipanthus* Hemandradenia* Jollydora* Manotes* Pseudoconnarus* Rourea...

 (1); Euphorbiaceae
Euphorbiaceae
Euphorbiaceae, the Spurge family are a large family of flowering plants with 300 genera and around 7,500 species. Most are herbs, but some, especially in the tropics, are also shrubs or trees. Some are succulent and resemble cacti....

 (1); Lecythidaceae
Lecythidaceae
The Lecythidaceae comprise a family of about 20 genera and 250-300 species of woody plants native to tropical South America and Madagascar.According to the most recent molecular analysis of Lecythidaceae The Lecythidaceae comprise a family of about 20 genera and 250-300 species of woody plants...

 (1); Maranthaceae (1); Melastomataceae
Melastomataceae
right|thumb|200px|Characteristic venation of many melastomesThe family Melastomataceae is a taxon of dicotyledonous flowering plants found mostly in the tropics comprising some 200 genera and 4500 species...

 (1); 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...

 (1); Myrtaceae
Myrtaceae
The Myrtaceae or Myrtle family are a family of dicotyledon plants, placed within the order Myrtales. Myrtle, clove, guava, feijoa, allspice, and eucalyptus belong here. All species are woody, with essential oils, and flower parts in multiples of four or five...

 (1); Anonaceae (1); Musaceae
Musaceae
Musaceae is a botanical name for a family of flowering plants. The family is native to the tropics of Africa and Asia. The plants have a large herbaceous growth habit with leaves with overlapping basal sheaths that form a pseudostem making some members appear to be woody trees.The family has been...

 (1); Apocynaceae
Apocynaceae
The Apocynaceae or dogbane family is a family of flowering plants that includes trees, shrubs, herbs, and lianas.Many species are tall trees found in tropical rainforests, and most are from the tropics and subtropics, but some grow in tropical dry, xeric environments. There are also perennial herbs...

 (1); Orchidaceae
Orchidaceae
The Orchidaceae, commonly referred to as the orchid family, is a morphologically diverse and widespread family of monocots in the order Asparagales. Along with the Asteraceae, it is one of the two largest families of flowering plants, with between 21,950 and 26,049 currently accepted species,...

 (1); Araceae
Araceae
Araceae are a family of monocotyledonous flowering plants in which flowers are borne on a type of inflorescence called a spadix. The spadix is usually accompanied by, and sometimes partially enclosed in, a spathe or leaf-like bract. Also known as the Arum family, members are often colloquially...

 (1); 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 ,...

 (1); Bignoriaceae (1).

In terms of importance value, palms are highly valued by all species of capuchin. In El Tuparro National Park in Colombia, the palm Attalea
Attalea (genus)
Attalea is a large genus of palms native to Mexico, the Caribbean, Central and South America. This pinnately leaved, non-spiny genus includes both small palms lacking an aboveground stem and large trees. The genus has a complicated taxonomic history, and has often been split into four or five...

 regiae
was a key species for this primate, the nuts being a principal food. In Manú National Park in Peru the palms Astrocaryum
Astrocaryum
Astrocaryum is a genus of about 36 to 40 species of palms native to Central and South America and Trinidad.-Description:Astrocaryum is a genus of spiny palms with pinnately compound leaves–rows of leaflets emerge on either side of the axis of the leaf in a feather-like or fern-like pattern. Some...

and Attalea were the most important palm genera, but perhaps not at the same level as Attalea in El Tuparro. Also, at Manú various species of Ficus
Ficus
Ficus is a genus of about 850 species of woody trees, shrubs, vines, epiphytes, and hemiepiphyte in the family Moraceae. Collectively known as fig trees or figs, they are native throughout the tropics with a few species extending into the semi-warm temperate zone. The Common Fig Ficus is a genus of...

were very important to the white-fronted capuchin; this emphasis on Ficus was not observed in the El Tuparro study, although this study did not include an entire year. Nevertheless, research on other species suggests the importance of palms as "key species" and the lack of importance of Ficus in such habitats as gallery forests in the llanos of Colombia and Venezuela, contrasts with their high importance in more fertile habitats like Manú.

The white-fronted capuchin takes advantage of almost any water source, drinking water from tree holes when available, but also drinking from brooks and springs when necessary. During the driest season in Vichada the group studied by Defler went to the ground every day to a water seep from under a huge boulder, which was the only water source available in their home range.

Reproduction

This species of capuchin is polygamous. The male mounts the female, holding her legs with his hind feet, and copulates with her for a few minutes. Although the time of 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 ....

 is unknown, it is probably around 160 days like the tufted capuchin. Usually one infant is born. Observations of a newborn in El Tuparro National Park showed the process by which the newborn discovered the appropriate position for riding on the mother. Newborns ride oriented sideways over the mothers' shoulders, but during the first days the baby holds on to any part of the mother such as the base of the tail, the tail, the legs, and the arms before discovering and learning that the position over the shoulders is best and most secure. After several weeks the baby makes the change from the sideways position over the shoulders to riding on her back.

All the members of the troop are interested in the newborn, and they take advantage of any opportunity to examine and look at its genitals if the mother permits it. With time the baby begins to climb up on other members of the troop, including the adult males who are interested in protecting the little one. Playing behavior is principally with a companion, and all members of the troop from the alpha male, the mother and all young members of the group solicit play with the young one.

Social structure

Adult males are notably tolerant of each other in the group, but they are very aggressive towards males of other groups. Intergroup aggressive behaviors have been observed in El Tuparro, which resulted in one group fleeing towards the central parts of their territory. Alpha males seem to exercise a "control position" at the center of the group, since all members are extremely conscious and alert to his location, and they all observe his reactions. If the alpha reacts with intense fear or panic or if he pays close attention to something, all members of the troop react similarly. The presence of adult males seems to lend psychological support to the smaller adult females. Defler noticed that more timid females often became quite aggressive towards him when a male appeared on the scene, although the females often needed to press up against the flank of the male for reassurance.

Communication

Vocalizations are variable, and some are listed as follows: (1) ua - a soft bark given repeatedly and used by all members of the group when danger is perceived; (2) ya - excited animals around the alpha, towards alpha and towards perceived danger; (3) eh-eh - threat towards potential danger, but especially adult females; accompanied with open mouth showing teeth (OMT); (4) squeaky hinge - threat given especially by young animals; (5) squeal - conflict within the group during a fight; (6) whistle - conflict in the group of a young animal; (7) ahr - a lost animal; others answer this call, apparently to direct it back to the group; (8) uh!uh!uh! - a common vocalization during feeding which may allow contact to be maintained and show general contment; (9) uch!uch! - an animal trying to keep up with the group; (10) warble - young animals establishing contact or coming close to an adult; (11) purr - close and pacific contact; (12) chirriar - pacific interaction of young ones during play.

Perhaps the most important display is the behavior of breaking branches, which all members of the group carry out. Even infants break small branches (or twigs), letting them fall to the soil, but the most spectacular is the alpha male who chooses large, dry branches which he hits with his hands and feet in spectacular jumps, so that they fall. Usually such branches make a tremendous sound as they fall through the other branches, and the members of the group become very excited and chatter loudly. This behavior is quite commonly discharged towards an observer when the animals have lost their fear.

Tool use

Trinidad white-fronted capuchins have been observed using leaves as cups to drink water from tree cavities. The leaves used were modified before by changing the shape of the leaf. The leaves are discarded after one use, meaning that a different leaf is used for repeat visits. These observations suggest that, like the common chimpanzee
Common Chimpanzee
The common chimpanzee , also known as the robust chimpanzee, is a great ape. Colloquially, the common chimpanzee is often called the chimpanzee , though technically this term refers to both species in the genus Pan: the common chimpanzee and the closely related bonobo, formerly called the pygmy...

, wild capuchins demonstrate tool manufacture and use in foraging-related contexts.

Interspecific interactions and predators

The white-fronted capuchin frequently travels with the common squirrel monkey
Common Squirrel Monkey
The common squirrel monkey is a small New World primate from the Cebidae family, and native to the tropical areas of South America.-Location:...

 and at other times with the tufted capuchin
Tufted Capuchin
The tufted capuchin , also known as brown capuchin or black-capped capuchin is a New World primate from South America...

 and Venezuelan red howler
Venezuelan Red Howler
The Venezuelan red howler is a South American species of howler monkey, a type of New World monkey. It is found in the western Amazon Basin in Venezuela, Colombia, Ecuador, Peru, Bolivia and Brazil...

. 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...

 often accompanies these monkeys, exactly as it does other species of primates. The white-fronted capuchin feels threatened by avian predators, and it is very vigilant around any large bird of prey. In Vichada, Colombia, Tayra
Tayra
The tayra , also known as the Tolomuco or Perico ligero in Central America, and San Hol or viejo de monte in the Yucatan Peninsula is an omnivorous animal from the weasel family Mustelidae. It is the only species in the genus Eira...

, Boa constrictor
Boa constrictor
The Boa constrictor is a large, heavy-bodied species of snake. It is a member of the family Boidae found in North, Central, and South America, as well as some islands in the Caribbean. A staple of private collections and public displays, its color pattern is highly variable yet distinctive...

and the Ornate Hawk-eagle have been seen trying to capture the monkey. After detecting the Tayra and Boa constrictor the members of the troop showed little fear and caution, even though these animals had been trying to capture the monkeys. The most common behavior after detecting a potential ground predator is "ya-ya" vocalization and branch breaking over the head of the potential predator, similar to the display of the brown woolly monkey
Brown Woolly Monkey
The brown woolly monkey or common woolly monkey or Humboldt's Woolly Monkey is a woolly monkey from Colombia, Ecuador, Peru and Brazil. It lives in groups of 2 to 70 individuals, usually splitting the group into smaller subgroups while in activity.Many published sources give the systematic name as L...

. In contrast, after being frightened by a male Ornate Hawk-eagle the monkeys screamed intensely only once, then hid quietly, some descending to the ground to sneak away.

Conservation status

The white-fronted capuchin is adaptable, has a wide distribution and probably is not yet endangered in Colombia on the species level. Nevertheless, it is probable that some subspecies are under considerable pressure. For example, C. a. cuscinus is found only is a small part of the SW Colombian Amazon and probably should be classified as "Vulnerable" for the country. We need to census the various subspecies and to clarify the taxonomy in order to evaluate properly the situation within the country. This species is found within 10–15 National Parks and it is probably not excessively hunted. Also, it survives well in secondary vegetation close to human beings.

External links

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