Tufted Capuchin
Encyclopedia
The tufted capuchin also known as brown capuchin or black-capped capuchin is a 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...

 from 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...

. As traditionally defined, it is one of the most widespread primates in the Neotropics, but it has recently been recommended considering the black-striped
Black-striped Capuchin
The black-striped capuchin, Cebus libidinosus, also known as the bearded capuchin, is a capuchin monkey from South America. It was the first non-ape primate where tool usage was documented in the wild, as individuals have been seen cracking nuts by placing them on a stone "anvil" while hitting them...

, black
Black Capuchin
The black capuchin, Cebus nigritus, also known as the black-horned capuchin, is a capuchin monkey from the Atlantic Forest in south-eastern Brazil and far north-eastern Argentina. Historically, it was included as a subspecies of the Tufted Capuchin....

 and Golden-bellied Capuchin
Golden-bellied Capuchin
The golden-bellied capuchin , also known as the yellow-breasted or buffy-headed capuchin, is one of several species of New World monkeys....

s as separate species, thereby effectively limiting the tufted capuchin to the Amazon Basin
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...

 and nearby regions.

The tufted capuchin is an omnivorous animal, mostly feeding on fruits and invertebrates, although it sometimes feeds on small vertebrates (e.g. lizards and bird chicks) and other plant parts. It can be found in many different kinds of environment, including moist tropical and subtropical forest, dry forest
Tropical and subtropical dry broadleaf forests
The tropical and subtropical dry broadleaf forest biome, also known as tropical dry forest, is located at tropical and subtropical latitudes. Though these forests occur in climates that are warm year-round, and may receive several hundred centimeters of rain per year, they have long dry seasons...

, and disturbed or 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...

.

Like other capuchins
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...

, it is a social animal
Social animal
A social animal is a loosely defined term for an organism that is highly interactive with other members of its species to the point of having a recognizable and distinct society.All mammals are social to the extent that mothers and offspring bond...

, forming groups of 8 to 15 individuals that are led by an alpha
Alpha (biology)
In social animals, the alpha is the individual in the community with the highest rank. Where one male and one female fulfill this role, they are referred to as the alpha pair...

 or dominant male.

Taxonomy and phylogeny

Previously all tufted capuchins were classified as Cebus apella. Under such taxonomy, the range of C. apella would extend throughout much of South America from Colombia to northern Argentina. More recent taxonomic studies have been carried out by Torres de Assumpção (1983; Torres 1988). Groves (2001, 2005) recognized the following subspecies for the tufted capuchins: C. apella apella (Linnaeus, 1758); C. apella fatuellus (Linnaeus, 1766); C. apella macrocephalus Spix, 1823; C. apella peruanus Thomas, 1901; C. apella tocantinus Lönnberg, 1939; C. apella margaritae Hollister, 1914; C. libidinosus libidinosus Spix, 1823; C. libidinosus pallidus Gray, 1866; C. libidinosus paraguayanus Fischer, 1829; C. libidinosus juruanus Lönnberg, 1939; C. nigritus nigritus (Goldfuss, 1809); C. nigritus robustus Kuhl, 1820; C. nigritus cucullatus Spix, 1823; C. xanthosternos Wied-Neuwied, 1826 (see Fragaszy et al. 2004; Rylands et al. 2005).

Physical characteristics

The tufted capuchin is more powerfully built than the other capuchins
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...

, with rougher fur and a short, thick tail. It has a bundle of long, hardened hair on the forehead that can be raised as a sort of "wig". The fur is brownish gray, with the belly being somewhat lighter-colored than the rest of the body. The hands and feet are black. The tail is strong and can be used as a grasping tail.

The tufted capuchin has a head-body length of 32 to 57 cm (12.6 to 22.4 in), a tail length of 38 to 56 cm (15 to 22 in), and a weight of 1.9 to 4.8 kg (4.2 to 10.6 lb), with the males generally being larger and heavier than the females.

Behaviour and ecology

The tufted 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...

, arboreal primate species, but it often forages on the ground to search for food or to walk longer distances between trees that are too far apart to jump.
The tufted capuchin lives in groups of two to twenty or more animals. A single group usually contains at least one adult male, but mixed groups with multiple males do also occur. In that case, one of the males is dominant. He accepts only a few monkeys in his direct surroundings, mainly younger animals and a few females. The dominant male and the group members that are close to him have the privilege to eat first in case of food scarcity, while subordinate monkeys have to wait until they are ready.

After a gestation period of 180 days, one young is born, or incidentally a twin. This young, which weighs only 200 to 250 grams, is carried on the back of its mother. The mother feeds her child for 9 months, but the young is sexually immature until its seventh year, which is quite late for a primate of its size.

Important natural enemies of the capuchin are large birds of prey
Bird of prey
Birds of prey are birds that hunt for food primarily on the wing, using their keen senses, especially vision. They are defined as birds that primarily hunt vertebrates, including other birds. Their talons and beaks tend to be relatively large, powerful and adapted for tearing and/or piercing flesh....

. They are so afraid of those birds, that they even become alarmed when a harmless bird flies over.

The tufted capuchin rubs urine
Urine
Urine is a typically sterile liquid by-product of the body that is secreted by the kidneys through a process called urination and excreted through the urethra. Cellular metabolism generates numerous by-products, many rich in nitrogen, that require elimination from the bloodstream...

 on its hands and feet in order to attract mates and reduce stress.

Diet

A recently discovered characteristic of one population this species is that it uses stones as a tool to open hard nuts. First it chooses ripe nuts from a nut palm. It uses its teeth to strip off the nut's fibrous husk. Then it leaves the nut to dry for about a week. When the nut is dry, the monkey lays the nut on a large, flat rock or fallen tree, hammering the nut with a suitable stone until the nut cracks. The hammer stones are often large enough to require lifting with both hands. The anvil rock is often pock-marked with hollows as a result of repeated use.

Besides nuts, the capuchin also eats fruit, insects, 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, egg
Egg (food)
Eggs are laid by females of many different species, including birds, reptiles, amphibians, and fish, and have probably been eaten by mankind for millennia. Bird and reptile eggs consist of a protective eggshell, albumen , and vitellus , contained within various thin membranes...

s, young birds, 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, and even bat
Bat
Bats are mammals of the order Chiroptera "hand" and pteron "wing") whose forelimbs form webbed wings, making them the only mammals naturally capable of true and sustained flight. By contrast, other mammals said to fly, such as flying squirrels, gliding possums, and colugos, glide rather than fly,...

s. They are also known to chase cats.

The tufted capuchin looks for its food in groups. As soon as one of the group members has found something edible, he or she may make a large whistling sound, dependent upon the proximity of other individuals and abundance of the food resource, so that the other monkeys know that there is something to eat. The composition of the group is very well organized, and is determined by rank in the hierarchy
Hierarchy
A hierarchy is an arrangement of items in which the items are represented as being "above," "below," or "at the same level as" one another...

. The dominant male often resides somewhere in the middle of the group just behind the front line, so that it is safer when a predator attacks. The vanguard is composed of higher-ranked females who are tolerated by the dominant male. They have the privilege to reach the food first, but they are also the most vulnerable when a predator attacks.

Tool use and manufacture

The tufted capuchin has been observed using containers to hold water, using sticks (to dig nuts, to dip for syrup, to catch ants, to reach food), using sponges to absorb juice, using stones as hammer and chisel
Chisel
A chisel is a tool with a characteristically shaped cutting edge of blade on its end, for carving or cutting a hard material such as wood, stone, or metal. The handle and blade of some types of chisel are made of metal or wood with a sharp edge in it.In use, the chisel is forced into the material...

 to penetrate a barrier and using stones as hammer
Hammer
A hammer is a tool meant to deliver an impact to an object. The most common uses are for driving nails, fitting parts, forging metal and breaking up objects. Hammers are often designed for a specific purpose, and vary widely in their shape and structure. The usual features are a handle and a head,...

 and 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...

 to crack nuts. While some of these tasks are relatively simple by cognitive standards (e.g. using a stick to catch ants), others, like cracking nuts with hammer and anvil are only exceeded in complexity by chimpanzees.

The potential for tool use in animals like the Tufted Capuchin depends on a number of conditions that would increase its likelihood of appearing in a given species. Van Schaik
Carel van Schaik
Carolus Philippus "Carel" van Schaik is a Dutch primatologist who since 2004 is professor and director of the Anthropological Institute and Museum at the University of Zürich, Switzerland....

 proposed that the occurrence of tool use would be likely in foraging species if three factors were present: manual dexterity, intelligence, and social tolerance. As it applies to manual dexterity, capuchins are capable of a limited precision grip (the ability to delicately pinch and manipulate objects with the thumb and fingertips), which is not found in any other New World monkeys and only found in limited amounts in apes. C. apella has an encephalization ratio
Encephalization quotient
Encephalization Quotient , or encephalization level is a measure of relative brain size defined as the ratio between actual brain mass and predicted brain mass for an animal of a given size, which is hypothesized to be a rough estimate of the intelligence of the animal.This is a more refined...

 greater than the hominids (except humans) and a neocortex ratio that is almost as large as the apes; both of these rough indicators suggest high intelligence. Finally, the Tufted Capuchin forms social groups typical of a complex and tolerant society.

The tufted capuchin has been observed manufacturing tools both in captivity and in the wild. In captivity, it has been reported as making probing sticks to reach normally inaccessible containers with syrup. It is also capable of understanding the concept of "sponging" and using paper towels, monkey biscuits, sticks, leaves and straw to sop up juice and then suck on the sponge to consume the juice. Research in the wild has shown that capuchin tool use is every bit as extensive as in captivity with capuchins being observed using stones to dig holes to get at tubers, an activity previously only seen in humans. The practice of using stones to crack nuts has arisen spontaneously in many locations such as in the Caatinga Dry Forest
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"...

 and Serra da Capivara National Park
Serra da Capivara National Park
Serra da Capivara National Park is a national park in the north east of Brazil. It has many prehistoric paintings. The park was created to protect the prehistoric artifacts and paintings found there. It became a World Heritage Site in 1991. Its head archaeologist is Niède Guidon...

, all in Brazil and hundreds of miles apart. It has been observed cracking various nuts and fruits such as palm nuts (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...

 and 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...

 spp.
) and jatobá fruits.(Hymenaea courbaril) The tufted capuchin has even been observed using stones to dislodge other stones that would later be used as hammers or shovels, an example of a more complex tool using behavior known as second-order tool use previously only found in chimpanzees. Curiously, not all tufted capuchins engage in tool use. Moura and Lee (2004) suggest lack of other food sources as the key factor. Ottoni and Mannu (2001), Fragaszy et al. (2004) and Visalberghi et al. (2005) have proposed this is likely more a factor of a monkey's terrestrial habit: the more time a monkey spends on the ground, the more likely it is to profit from (and thus engage in) tool use.

In captivity, the tufted capuchin has been seen to manufacture stone tools that produced simple flakes
Lithic flake
In archaeology, a lithic flake is a "portion of rock removed from an objective piece by percussion or pressure," and may also be referred to as a chip or spall, or collectively as debitage. The objective piece, or the rock being reduced by the removal of flakes, is known as a core. Once the proper...

 and cores
Lithic core
In archaeology, a lithic core is a distinctive artifact that results from the practice of lithic reduction. In this sense, a core is the scarred nucleus resulting from the detachment of one or more flakes from a lump of source material or tool stone, usually by using a hard hammer percussor such...

. Some of the capuchins even used these sharpened stones to cut (in a back-and-forth motion) barriers in order to reach food. The importance of this behavior is that it serves as evidence of mechanical proclivity to modify stones by using behaviors already in the monkeys' repertoires, and this behavior is seen as a precursor to stone-knapping
Lithic reduction
Lithic reduction involves the use of a hard hammer precursor, such as a hammerstone, a soft hammer fabricator , or a wood or antler punch to detach lithic flakes from a lump of tool stone called a lithic core . As flakes are detached in sequence, the original mass of stone is reduced; hence the...

. This early and limited tool use behavior has been hypothesized as similar to pre-Homo habilis
Homo habilis
Homo habilis is a species of the genus Homo, which lived from approximately at the beginning of the Pleistocene period. The discovery and description of this species is credited to both Mary and Louis Leakey, who found fossils in Tanzania, East Africa, between 1962 and 1964. Homo habilis Homo...

and that artifacts of that time would probably resemble those of capuchins.

C. apella tool manufacture and use has been analyzed for potential clues to social learning
Observational learning
Observational learning is a type of learning that occurs as a function of observing, retaining and replicating novel behavior executed by others...

 and problem solving ability, as tool manufacture and use can often shed light on such complex cognitive abilities. Social learning, or the ability to learn from other individuals, is a controversial topic in most nonhuman species like C. apella because of the relative difficulty of determining whether a behavior was learned from imitation
Cognitive imitation
Cognitive imitation is a type of imitation and a type of social learning. Cognitive imitation, like the imitation of motor rules , involves learning and copying specific rules by observation. The principal difference between motor and cognitive imitation is the type of rule that is learned and...

 or a much simpler form of social learning. One way of closing the gap between concurrent tool related behaviors and their likelihood of arising from imitation is by narrowing down events that would make social learning more probable such as a preference for observing experienced tool users. In this regard, Ottoni and his team found that young capuchins tended to observe the best tool users when cracking nuts. Another way of isolating imitation from other simpler behaviors is to present the capuchins with a box that has food but has two different ways of opening it. The important point is that neither way should be more advantageous so that the monkey can freely choose one. In one such study, when humans opened the door in front of the monkeys using one way only, the monkeys used that method, even when they discovered the alternative on their own. In another study, capuchin alphas from two separate groups were trained to open the door in a specific way, after which the monkeys were paired with subordinates who learned to open the door in the same way. When capuchins are trained in the same way and this time released into their groups, the habit is once again disseminated amongst all group members even when others discover alternative ways. Nevertheless, the subject of whether or not C. apella learns by imitation is still controversial, because of the inherent difficulty in teasing out unambiguous evidence of a complex cognitive process such as imitation.

Problem solving

Tool use and manufacture can also shed light on the many aspects of the tufted capuchin's cognitive abilities by determining how it solves some problems. Some non-primates manufacture and use objects as tools. Crows are known to make hook-tools for catching insects, but such activities lack the behavioral plasticity of tool use as evidenced in tufted capuchins who found new ways to use tools that other species could not. But this plasticity in tool use, while suggesting greater complexity and cognitive ability, does not suggest that the monkeys understand cause and effect. It instead implies they are only able to learn from successful efforts but not from failures nor are they able to refine and improve much. Its ability to repeat successes, coupled with its complex repertoire of behavioral events helps to explain the Tufted Capuchin's extensive repertoire of innovative behaviors besides tool use.

Distribution and habitat

This species lives in the northern Amazon rainforest
Amazon Rainforest
The Amazon Rainforest , also known in English as Amazonia or the Amazon Jungle, is a moist broadleaf forest that covers most of the Amazon Basin of South America...

 of the Guyanas
Guyanas
Guyana is a country in South America.Guyana, Guayana, or Guiana may also refer to:*Guayana Esequiba, the territory of Guyana claimed by Venezuela*Guayana Region, an administrative region of Venezuela...

, Venezuela and 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...

 and to the west of the Rio Negro, as far north as the Orinoco
Orinoco
The Orinoco is one of the longest rivers in South America at . Its drainage basin, sometimes called the Orinoquia, covers , with 76.3% of it in Venezuela and the remainder in Colombia...

 in 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...

. It is also found in eastern Colombia, Ecuador, Peru, including the upper Andean Magdalena valley in Colombia. It can be found in a large variety of forest types, mainly in tropical rainforest
Tropical rainforest
A tropical rainforest is an ecosystem type that occurs roughly within the latitudes 28 degrees north or south of the equator . This ecosystem experiences high average temperatures and a significant amount of rainfall...

s (up till a height of 2700 m), but also in more open forests.

The distribution overlaps with that of other species of capuchins, such as 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...

(Cebus albifrons).

External links

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