BLACK-CAPPED CAPUCHIN

Sapajus apella

Geographic Distribution and Habitat
Black-capped capuchins (Sapajus apella), also known as tufted capuchins, are native to South America, where they are ubiquitous throughout the Amazon River Basin. Extant populations reside in the countries of Bolivia, Brazil, Colombia, Ecuador, French Guiana, Guyana, Peru, Suriname, and Venezuela. A breeding population was introduced to the Caribbean neotropics. These monkeys reside on the northwestern peninsula (Chaguaramas) of Trinidad Island within the Republic of Trinidad and Tobago, close to the South American continent, northeast of Venezuela and northwest of Guyana. They share the island with two endemic monkey species: the white-fronted capuchin (Cebus albifrons trinitatis) and the Trinidad red howler monkey (Alouatta seniculus insulanus). The only other Caribbean island to host neotropical primates is Isla de Margarita, home to the Margarita Island capuchin (Sapajus apella margaritae). 

A variety of forest types cover the monkeys’ vast range, including: tropical, subtropical, dry, submontane, savannah, mangrove, disturbed or secondary, and várzea (seasonally inundated) forests. Black-capped capuchins dwell in all these habitats. They have been sighted in tropical rainforests at altitudes up to 8,858 ft (2,700 m) and also in tropical lowlands and open forests. The species is most comfortable, however, within the relative safety of the dense tree canopy (particularly, the lower and middle levels), which provides shelter, sleeping quarters, food sources, and ease of travel.

The Caribbean black-capped capuchins live on habitat islands surrounded by towns, agricultural areas, and roads.

TAXONOMY IN TRANSITION

The International Union for Conservation of Nature (IUCN) revealed in its July 2020 updates of the IUCN’s Red List of Threatened Species that evolutionary analyses of the Sapajus genus conducted in 2017 and 2018 lumped several previously considered species together as a single, widespread lineage. Those previously considered separate capuchin species are the Azaras’s capuchin (S. cay), the bearded capuchin (S. libidinosus), the tufted capuchin (S. apella), the large-headed capuchin (S. macrocephalus), and the blond capuchin (S. flavius).

It is important to note that, for conservation purposes, the IUCN has opted to treat separately the Azaras’s capuchin (S. cay) (ranked Least Concern), the bearded capuchin (S. libidinosus) (Vulnerable), and the blond capuchin (S. flavius) (Critically Endangered). The justification is that each bears distinct external characteristics, occurs in regions with distinct flora and fauna, occupies distinct adaptive zones, and faces particular conservation issues and threats.

However, the large-headed capuchin is now considered to be synonymous with the tufted capuchin (S. apella).

With updated taxonomy also comes parenthood—or demotion, depending on one’s perspective. The IUCN has deemed the black-capped (tufted) capuchin as the “parent” species to both the Guianan brown capuchin (Sapajus apella apella) and the Margarita Island capuchin (Sapajus apella ssp. margaritae). Before the IUCN relegated these two monkeys to subspecies ranking, each had been regarded as an individual species.

Black-capped capuchin range, IUCN, 2020

Size, Weight, and Lifespan
Black-capped capuchin males are typically larger than the females, an example of sexual dimorphism in the species. Average weight for an adult male black-capped capuchin is 6.7 lb (3 kg), with a range of 3–10.5 lb (1.35–4.8 kg). Average weight for an adult female is 5.3 lb (2.4 kg) with a range of 4–7.5 lb (1.76–3.4 kg). Captive black-capped capuchins tend to be heavier than those in the wild. A more sedentary lifestyle, and perhaps a deficient diet, might explain the extra pounds. Males are toothier than their female counterparts, with canines that are 22 percent larger than those of the ladies.

Head-to-body length for males is about 17 in (44 cm) and 15 in (39 cm) for females. The monkeys’ tails nearly equal their bodies in length.

In the wild, average lifespan for this species is between 15 and 20 years. Life as a captive may extend a black-capped capuchin’s existence to nearly 50 years.

Appearance
Black-capped capuchins are sturdy New World monkeys who look like they might spend a lot of time at the gym—the jungle gym, of course (what were you thinking?). Their robust physique distinguishes them from the more delicate gracile capuchins in the Cebus genus. A thick rug of a fur coat (pelage) covers their powerful frame. Shoulders, upper arms, and torso are varying shades of light brown to honey mustard with intermingling strands of gray. Black fur adorns the remainder of the body, including the monkey’s long, thick tail. Hands and feet are dark brown to black.

But it’s the remarkable hairdo that gives this primate its common name of “black-capped capuchin” and secondary name of “tufted capuchin.” A black cap of coarse fur that, in some individuals, looks like a failed mohawk sits atop the crown of the head in unruly tufts that tickle the top of the ears. White fur sometimes outlines the cap, toward the face, creating a “V” boundary.

A pensive face with pale, pinkish skin is framed by long black sideburns that extend from the cap downward and get lost beneath the chin. Brown eyes assess the world from their perch above a broad, flat muzzle fitted with inquiring nostrils.

A Tufted Capuchin runs on its hind legs and tries to break a coconut in Rio de Janeiro Botanical Gardens

Diet
Lots of fruits, along with seeds, insects, frogs, lizards, squirrels, bats, birds, eggs, the occasional small mammal (such as the mouse opossum), carrion, and a sprinkling of stems, flowers, and leaves comprise the diet of black-capped capuchins. Nature has fitted these omnivores with a deep lower jaw and powerful jaw muscles that easily accommodate larger fruits and rough vegetation. Pith from the Scheelea palm tree provides critical sustenance to the monkeys during the dry season, when food is scarce.

A field study from 2005 reports on, what is believed to be, an isolated incident of an infant red-bellied titi monkey (Callicebus moloch) falling prey to hungry black-capped capuchins. Another wildlife study published in 2008 reports on a group of black-capped capuchins feeding on a deceased adult female owl monkey (Aotus brumbacki) in a fragment of gallery forest in the Colombian Llanos. The witnessing field scientists determined that the monkeys appeared to be scavenging the owl monkey’s carcass, rather than having killed the animal. Nevertheless, they deemed the event remarkable for the capuchins’ potential to exploit potential prey of this size.

Behavior and Lifestyle
Black-capped capuchins are predominantly arboreal (spending most their time in trees), moving quadrupedally (on all four limbs) by leaping and climbing through the trees. They sometimes descend to the ground for play or to cross areas of scarce vegetation; they are able to walk upright (bipedally) for short distances. A prehensile tail aides in seizing and grasping, but it only supports the full body weight of juvenile (and lighter-weight) monkeys, who are able to hang fully suspended from a tree branch by their tail. Adult (and heavier) monkeys may also use their tail to hang from a branch but need the help of another limb to keep from falling. Compared to other New World monkeys, the tail of the black-capped capuchin is less flexible and completely covered by fur. Black-capped capuchins are the only species of capuchin monkey who, while moving through habitat, carry their tails in a tight coil.

A diurnal species (active during daylight hours), the monkeys spend most of their day, except for a midday nap, foraging and eating. Overnight, they sleep in the trees, tucked between branches.

These funny-hairdo monkeys have a reputation as noisy and destructive foragers. Their reputation is deserved. As they advance through the forest, leaping from tree to tree, they wantonly rip apart vegetation in their path. To crack open nuts, they slam the encased fruits against tree branches. With their large teeth, they tear into large fruits that have coriaceous (leathery) skins and loudly chew with assistance from their formidable jaws. In their search of vertebrate and invertebrate prey, the monkeys disrupt dead vegetation, capturing and eating the small creatures they find in these nests. They are not likely to share caught prey with members of their troop.

Whatever black-capped capuchins might lack in dining etiquette, they make up for with their impressive manual dexterity, precision, and problem-solving abilities. While other monkey species are left scratching their heads (figuratively speaking), black-capped capuchins successfully exploit tough-to-get-at food resources, deftly extracting fruits and prey.

They are master tool users and innovators. In their capable hands, long sticks become probes to extract insects from holes. Stones become multiple instruments with multiple uses: a hammer, wielded by the monkeys with precision and great force to crack open hard fruits and nuts placed upon a large flat stone serving as makeshift anvil; a hammer and chisel to break through barriers; or a shovel to dig up tubers. The monkeys use forest sponges to absorb fruit juices and syrup, and they fashion natural containers to hold their water reserves. The species’ prowess has been found only in chimpanzees (and perhaps in some exceptional human primates).

Scientists postulate that black-capped capuchins create “mental maps,” enabling them to locate nourishing foods while foraging efficiently and with optimum energy. While the monkeys’ excellent posture control, strong core, and long limbs enable them to physically perform, scientists credit the monkeys’ problem-solving capabilities to brain size. With respect to their body weight, black-capped capuchins are fitted with larger-than-expected brains.

As you might imagine, rainforests are home to a denizen of insects—biting, annoying insects if you are a mammal, like the black-capped capuchin. But these resourceful monkeys have discovered a method to repel insects and parasites. In a practice scientifically known as “anointing,” or more casually, “anting”—or “fur rubbing”—they crush acidic ants or millipedes and then rub the squashed creatures, mixed with the monkeys’ own urine, against their bodies. Mosquitoes and ticks are dissuaded by the offensive odor and seek out another warm-blooded host to feast upon. Including urine in the pungent mix may help to regulate the monkey’s body temperature, may reduce stress, and might even be a “turn on” for attracting a mate. Males are known to rub urine between their hands, on their feet, and on the tip of their tail to convey their sexual virility to females.

Daily Life and Group Dynamics
Black-capped capuchins are playful, curious, intelligent monkeys who live in family groups (known as “troops”) of 15 to 20 individuals, though smaller groups are known to occur. Troop composition includes one or more adult males, several adult females with young offspring, and juveniles. The dominant male, known as the “alpha,” is the troop’s leader and elder. He protects members of his troop from predators and from rival monkey troops (aggression with outside troops is most often over food resources) and takes the lead in defending his troop’s territory. The alpha male also directs his troop’s foraging expeditions, decides when individuals can take a rest, and chooses sleeping sites for the night. His rank gives him certain privileges: when it comes to food, he gets first choice and eats first. Likewise, he gets first choice of his “harem” when it comes to mating. His main female partner, the dominant female of the troop, is ranked second-in-command in hierarchy. The troop’s social composition ensures that the genes of the more dominant and fit monkeys are passed on to the next generation.

As they grow into adulthood, a troop’s subordinate males usually leave their birth (natal) group in search of joining a troop where they might rise to the rank of alpha male. Females, however, spend their entire lives with their birth group.

Home ranges are between 62 and 99 acres (25–40 ha), but can be as wide as 877 acres (355 ha); typical day range for foraging is about ½ acre (2,000 m).

Social composition of a troop’s foraging expedition is well organized, determined by hierarchal rank, with the alpha male and most dominant female occupying the central-forward positions and other dominant females at the front line. When a troop member discovers a food source, she alerts the other monkeys with a loud whistle. The more dominant members get first pick, while lower-ranking individuals get leftovers. To ensure they get a decent bite to eat (especially when food is scarce), subordinate monkeys have been known to “fake out” dominant monkeys by sounding an alarm call—instead of a food whistle. Hearing the alarm call, and believing that a predator is nearby, the dominant monkeys flee, leaving the lower-ranking individuals to a usurped meal.

Black-capped capuchins have been observed foraging with squirrel monkeys (genus Saimiri). It’s a cooperation that seems of more benefit to the squirrel monkeys. Because the black-capped capuchins are so enthusiastic in tearing through vegetation, squirrel monkeys are able to locate food much quicker than foraging alone. Also, should a capuchin find a fabulous patch of fruit, he whistles to the others in his troop to join the feast. Squirrel monkeys might be uninvited banquet guests.

Geographic distribution of black-capped capuchins overlaps with that of other capuchin species, including the white-fronted capuchin.

Large raptors are the natural predators of black-capped capuchins. So inherent is their fear of these birds that even when a harmless bird flies overhead, the monkeys become agitated and sound alarm calls.

What Does It Mean?

Alarm call:
Specific calls that individuals in a troop make to warn other members of their group of imminent danger, such as predators.

Biome:
A large naturally occurring community of flora and fauna occupying a major habitat, e.g. forest or tundra.

Canopy:
Beneath the emergent layer, the canopy layer is the primary layer of the forest and forms a roof over the two remaining layers (the understory and forest floor). Many animals live in this maze of leaves and branches, where food is abundant.

Carrion:
The decaying flesh of dead animals.

Contact call:
Conveys an individual’s location to other members of a group while foraging for food.

Disturbed secondary forest:
Disturbances refer to events that upset a habitat resulting in substantial impact. Disturbances include windstorms, logging, fire, and floods. A secondary forest is a forest that has re-grown after a major disturbance, such as fire or timber harvest, but has not yet reached the mature state of primary forest.

Dominance hierarchy:
A type of social hierarchy that arises when members of a social group interact, often aggressively, to create a ranking system. In social living groups, members are likely to compete for access to limited resources and mating opportunities.

Estrus:
A recurring period of sexual receptivity and fertility in many female mammals.

Extant:
Currently or already existing or living.

Extractive foraging: 
The act of finding and processing food sources found embedded in the ground, sometimes with the aid of tools.

Flagship species:
A species selected to act as an ambassador, icon, or symbol for a defined habitat, issue, campaign or environmental cause.

Gallery forest: 
A forest that serves as a corridor along rivers or wetlands and projects into landscapes that are otherwise sparsely treed, such as savannas, grasslands, or deserts.

Habitat fragmentation:
The process by which a large, continuous stretch of habitat gets divided into smaller, disjointed patches of habitat.

Harem:
A group of females guarded by a male, who prevents other males from mating with them; an animal group consisting of one or two males, a number of females, and their offspring. The dominant male drives off other males and maintains the unity of the group. If present, a second male is subservient to the dominant male.

Homology:
The state of having the same or similar relation, relative position, or structure.

Lowland forest:
An equatorial evergreen rainforest, commonly known as a tropical rainforest, which receives high rainfall (80 in; 2 m) throughout the year.

Mangrove: 
A mangrove refers to two different things: a tidal swamp ecosystem found in tropical deltas, estuaries, lagoons, or islands, and the characteristic tree species populating this ecosystem. Mangrove trees have developed unique adaptations to the harsh conditions of coastal environments.

Neotropics:
Span from central Mexico and the Caribbean islands to southern South America. The Neotropical Region contains the largest number of mammal species in the New World, the second-highest number of mammal species on the planet, and is the region with the largest number of endemic mammal families in the world.

Omnivore:
An animal or person who eats food of both plant and animal origin.

Pith:
The soft or spongy tissue of a plant or fruit, which is usually white or pale in color (i.e., the white part between the skin and fruit of an orange). 

Polygamous:
Having more than one mate.

Prehensile tail: 
The tail of an animal that has adapted to be able to grasp or hold objects.

Savanna forest:
A mixed woodland grassland ecosystem characterized by the trees being widely spaced so that the canopy does not close. Usually found in tropical or subtropical regions with seasonal rainfall.

Secondary forest:
A forest that has regrown after a major disturbance, such as fire or timber harvest, but has not yet reached the mature state of primary forest.

Sexual dimorphism:
Distinct difference in size or appearance between the sexes of an animal in addition to difference between the reproductive organs themselves.

Submontane forest:
A forest that is situated at the foothills or lower elevation slopes of a mountainous region.

Subspecies:
A population usually restricted to a geographical area that differs from other populations of the same species, but not to the extent of being classified as a separate species.

Subtropical forest:
A type of forest found in areas outside of the strict equatorial region—but either within or bordering the tropical zone—and have more noticeable seasonal changes. Although temperatures may vary only slightly over a year, rainfall may be distributed unevenly so that wet and dry seasons occur.

Sympatric: 
Occurring or living in the same area; overlapping in distribution.   

Taxonomy: 
A branch of science that encompasses the description, identification, nomenclature, and classification of organisms.

Tropical forest:
A type of forest found in areas with high regular rainfall and no more than two months of low rainfall, and consisting of a completely closed canopy of trees that prevents penetration of sunlight to the ground and discourages ground-cover growth.

Várzea forests:
A seasonal floodplain forest inundated by whitewater rivers that occurs in the Amazon biome.

Vertebrates:
Animals with a backbone, including mammals, birds, reptiles, amphibians, and fish.

Visit the Glossary for more definitions

Communication
Black-capped capuchin monkeys communicate with one another through vocalizations, body language, tactile methods, and olfactory cues.

Vocalizations include contact calls, calls emitted at the end of aggressive encounters, alarm calls, whistles sounded while foraging, and shrieks emitted during sexual encounters. Facial expressions, gestures, and postures convey mood or intention. Mutual grooming sessions are important in cultivating social bonds with one another. Playtime, besides being fun, teaches young capuchins social boundaries.

Some scientists conjecture that the practice of anointing, or “anting” (referring to the monkeys’ use of crushed ants as insect repellent), helps to strengthen social bonds. This conjecture may be true of other capuchin species, who practice “social anointing”—whereby the monkeys anoint one another to improve coverage of topically applied insect and anti-parasite repellants. However, black-capped capuchins rarely engage in this practice, instead sticking to individual anointing (or anting). To illuminate: individual anointing, practiced on oneself (and keeping one’s hands to oneself), targets “hard-to-see” body parts that are difficult to groom. Social anointing, practiced by multiple individuals on one another, is regarded as a mutual medication session, targeting “hard-to-reach” body parts on another.

Other scientists hypothesize that anointing creates a “group scent,” leaving important olfactory cues. But whatever cue or message the anointing capuchins intend is a mystery to scientists—unlike, for example, the cue that says, “Let’s get it on,” sent by randy males to females when the males rub their own urine on their hands, feet, and tail.

Regardless of the monkeys’ intention of their olfactory proclivities, scientists agree that black-capped capuchins possess strongly developed olfactory communication.

Fun Facts

Capuchins are named after friars (belonging to the order of Franciscan monks) who dressed in brown robes with hoods covering their heads.

As of 2012, capuchins are divided into two genera: gracile capuchins, in the genus Cebus, and robust capuchins, in the genus Sapajus. Robust capuchins are, as the name implies, more sturdily built than gracile capuchins. Their limbs are proportionally shorter, and their canine teeth are shorter and thicker.

Reproduction and Family
Females attain sexual maturity (able to conceive and bear young) between 4 to 6 years of age. Males lag behind a bit, attaining sexual maturity (able to sire young) at 7 years of age.

The species is polygamous. A troop’s females mate mostly with the alpha male and do their best to attract his attention, eschewing a troop’s subordinate males. If the alpha male is not present, subordinate males might get lucky, however.

When she comes into estrus and wishes to mate, a female calls out loudly to the male, trailing him wherever he goes. She does her best to solicit him through flirtations, raising her eyebrows and giving him an exaggerated smile. She might reach out and touch the male before quickly running away, enticing him to follow. But the male isn’t so easily snagged. He plays hard to get and ignores the aroused female, forcing her to vie for his attention with further antics (she might throw rocks at him to get his attention). Eventually, the male relents and permits the determined female to sit beside him. Soon after, the two copulate. Both partners emit vocalizations during the sex act.

Black-capped capuchins mate year-round, though most births tend to occur during the spring and summer months (the dry to early rainy seasons), when there is plenty of fruit to eat.

After a pregnancy of about 155 days, a single infant is born. Twin births are rare. A mother nurses her baby for 9 months and provides most the parental care. For the first several months of her life, a baby clings to her mother as mom travels through the forest.

Ecological Role
Black-capped capuchins help regenerate their forest habitat by dispersing seeds (through their feces) from the great variety of fruits they eat. They also help preserve their ecosystem by eating insects that would otherwise destroy trees. Lastly, their practice of “pruning” encourages the trees where they roost to regenerate more branches that bare additional fruit.

Conservation Status and Threats
Due to the species widespread geographic distribution, no immediate threats to its survival, and a high reproductive potential, the black-capped capuchin is classified as Least Concern by the International Union for Conservation of Nature (IUCN, 2020), appearing on the IUCN Red List of Threatened Species. The overall population remains stable, although the species is declining in some areas of its range due to hunting and habitat loss.

Monkeys are hunted and killed for their flesh, known as “bushmeat.” Infant black-capped capuchins, left helpless when their mothers are murdered, are kidnapped to be sold as pets in the illicit wildlife trade. Increasingly, natural habitat is razed to accommodate human developments, including roadways and settlements. Coffee and eucalyptus plantations replace forests.

And while they may not be the favored capuchin for biomedical research (the white-throated capuchin, Cebus capucinus, also known as the white-faced capuchin, might hold this indignity), black-capped capuchins are also unwitting subjects, held captive in nonhuman primate laboratories. Citing the monkeys’ “anatomic and physiologic similarities and genetic homology with humans,” scientists have experimented on black-capped capuchins to study abnormal heart rhythms and cardiovascular diseases. They’ve also exploited the monkeys to study the field of dentistry.

Other captive black-capped capuchins are exploited by Hollywood, forced to become actors in movies and television shows. Capuchin monkeys—white-throated capuchins, in particular—have been forced to entertain humans in other ways: as sidekick to the “organ grinder,” an arcane human street performer; and as monkey jockeys, strapped onto the backs of greyhound dogs in the early days of greyhound racing (a dying industry that, on its own, is rife with cruelty and abuse). More recent times have seen these capuchins forced to ride sheepdogs in “monkey rodeos.”

Like other capuchin species, black-capped capuchins have been used as service animals, trained to perform simple tasks for humans with limiting physical disabilities. But in March 2011, the Americans with Disabilities Act (ADA) revised its definition for service animal. Citing animal welfare concerns, the potential for serious injury to people, and risks of transferring dangerous diseases to humans, the ADA removed nonhuman primates from its grace. Today, the ADA recognizes only dogs as service animals. It must be noted, however, that some state and local laws define “service animal” more broadly than the ADA’s definition. Adding to this legislative murkiness is the legality or illegality of keeping capuchin monkey as pets, depending on state (in the U.S.).

Conservation Efforts
The black-capped capuchin is listed in Appendix II of the Convention on International Trade in Endangered Species (CITES), an international agreement between governments whose goal is to ensure that international trade in specimens of wild animals and plants does not threaten their survival.

Numerous national parks and wildlife reserves throughout the countries of the species’ vast range provide protected habitat to those monkeys who reside within the boundaries.

Neotropical Primate Conservation (NPC) is a wildlife conservation group, founded in the United Kingdom in 2007 with branches in Peru (2014) and Colombia (2016), dedicated to the conservation of primates and their habitats in South and Central America. Black-capped capuchins fall under the auspices of NPC. The group promotes conservation and protects biodiversity in the neotropics through land protection, research, improved degraded habitat for wildlife, public awareness/educational campaigns. and facilitating the livelihoods of local people by helping them to create sustainable, ecological products. Since 2007 NPC has been using primates as “flagship species” for its community conservation project.

The Wild Capuchin Foundation is a California (USA)-based nonprofit, founded in 2012, dedicated to the conservation of capuchin monkey habitat. Its main focus is on the white-faced capuchin (Cebus capucinus) (aka, white-throated) population of Lomas Barbudal, Costa Rica. The foundation is a partner to the Lomas Barbudal Monkey Project, supporting the project’s scientific, educational, and conservation work.

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Written by Kathleen Downey, February 2021