Tufted capuchin [1] | |
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Primates |
Suborder: | Haplorhini |
Infraorder: | Simiiformes |
Family: | Cebidae |
Genus: | Sapajus |
Species: | S. apella |
Binomial name | |
Sapajus apella | |
Subspecies | |
Sapajus apella macrocephalus Sapajus apella apella | |
Geographic range following taxonomy used by IUCN | |
Range of subspecies macrocephalus | |
Synonyms | |
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The tufted capuchin (Sapajus apella), also known as brown capuchin, black-capped capuchin, or pin monkey, is a New World primate from South America and the Caribbean islands of Trinidad and Margarita. 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 and golden-bellied capuchins as separate species in a new genus, thereby effectively limiting the tufted capuchin to the Amazon basin and nearby regions. [1] However, the large-headed capuchin (S. a. macrocephalus), previously defined as a distinct species, has been reclassified as a subspecies of the tufted capuchin, expanding its range east to Peru and Ecuador and south to Bolivia. [3]
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, and disturbed or secondary forest.
Like other capuchins, it is a social animal, forming groups of 8 to 15 individuals that are led by an alpha or dominant male.
At one point all tufted capuchins were classified as Cebus apella. [4] [5] Under such taxonomy, the range of C. apella would extend throughout much of South America from Colombia to northern Argentina. Although she did not describe specific or subspecific nomenclature, Torres de Assumpção (1983; 1988) described differences between tufted capuchins from five distinct geographic regions of Brazil and high phenotypic variation of individuals in a sixth area that had a greater selection pressure. [6] [7] In 2001, Silva Júnior proposed that the robust capuchins such (formerly the C. apella group) be placed in a separate genus, Sapajus, from the gracile capuchins (formerly the C. capucinus group) which retain the genus Cebus. [8] This was supported by Jessica Lynch Alfaro et al. in 2011. [9] [10] Groves (2005) recognized six subspecies: Cebus apella apella, C. a. fatuellus, C. a. macrocephalus , C. a. margaritae, C. a. peruanus, C. a. tocantinus. [1] The IUCN follows Silva (2001) and recognise the species as monotypic, though the subspecies status of S. a. margaritae is unclear. [2]
The tufted capuchin is more powerfully built than the other capuchins, with rougher fur and a long, 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 prehensile: strong and can be used for grasping, as an extra limb.
The tufted capuchin has a head-body length of 32 to 57 centimetres (13 to 22 in), a tail length of 38 to 56 centimetres (15 to 22 in), and a weight of 1.9 to 4.8 kilograms (4.2 to 10.6 lb), with the males generally being larger and heavier than the females.
The tufted capuchin is a diurnal, 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 (7.1 to 8.8 oz), is carried on the back of its mother. The mother feeds her child for 9 months, but the young are 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. They are so afraid of those birds that they even become alarmed when a harmless bird flies over.
A recently discovered characteristic of one population of 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, [11] hammering the nut with a suitable stone until the nut cracks. The hammerstones are often large enough to require lifting with both hands. The anvil rock is often pockmarked with hollows as a result of repeated use. [12] [13]
Besides nuts, the capuchin also eats fruit, leaves, [14] seeds, pith, [15] insects and larvae, eggs and young birds, frogs, lizards, other reptiles, [15] rodents, mouse opossums, [15] and even bats. 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. [16] The composition of the group is very well organized and is determined by rank in the hierarchy. 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. It is a host of the acanthocephalan intestinal parasite Pachysentis rugosus . [17]
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 to penetrate a barrier [18] and using stones as hammer and anvil to crack nuts. [19] 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. [20]
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 proposed that the occurrence of tool use would be likely in foraging species if three factors were present: manual dexterity, intelligence, and social tolerance. [21] 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. [22] [23] C. apella has an encephalization ratio 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. [24] [25] Finally, the tufted capuchin forms social groups typical of a complex and tolerant society. [26] [27]
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. [28] 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. [28] 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. [29] The practice of using stones to crack nuts has arisen spontaneously in many locations such as in the Caatinga Dry Forest [29] and Serra da Capivara National Park, [30] all in Brazil and hundreds of miles apart. It has been observed cracking various nuts and fruits such as palm nuts ( Attalea and Astrocaryum spp.) [31] and jatobá fruits ( Hymenaea courbaril ). [30] 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. [30] Curiously, not all tufted capuchins engage in tool use. Moura and Lee (2004) [32] suggest lack of other food sources as the key factor. Ottoni and Mannu (2001), [33] Fragaszy et al. (2004) [31] and Visalberghi et al. (2005) [34] 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 and cores. Some of the capuchins even used these sharpened stones to cut (in a back-and-forth motion) barriers in order to reach food. [35] 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. [22] This early and limited tool use behavior has been hypothesized as similar to pre- Homo habilis and that artifacts of that time would probably resemble those of capuchins. [35]
S. apella tool manufacture and use has been analyzed for potential clues to social learning and problem solving ability, as tool manufacture and use can often shed light on such complex cognitive abilities. [18] [36] Social learning, or the ability to learn from other individuals, is a controversial topic in most nonhuman species like S. apella because of the relative difficulty of determining whether a behavior was learned from imitation or a much simpler form of social learning. [37] 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. [19]
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. [38] 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. [39] 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. [40] Nevertheless, the subject of whether or not S. 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. [38] [41]
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, [42] 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. [43] 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. [44] 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. [44]
This species lives in the northern Amazon rainforest of the Guyanas, Venezuela and Brazil and to the west of the Rio Negro, as far north as the Orinoco in Venezuela. It is also found in eastern Colombia, Ecuador, Bolivia, Peru, including the upper Andean Magdalena valley in Colombia. An introduced breeding population is well established in the northwestern peninsula of the island of Trinidad in the Republic of Trinidad and Tobago. The subspecies/population on Margarita Island in Venezuela, S. a. margaritae, is considered Critically Endangered by the IUCN Red List. [45] It can be found in a large variety of forest types, mainly in tropical rainforests (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 (Cebus albifrons).
The Cebidae are one of the five families of New World monkeys now recognised. Extant members are the capuchin and squirrel monkeys. These species are found throughout tropical and subtropical South and Central America.
The Panamanian white-faced capuchin, also known as the Panamanian white-headed capuchin or Central American white-faced capuchin, is a medium-sized New World monkey of the family Cebidae, subfamily Cebinae. Native to the forests of Central America, the white-faced capuchin is important to rainforest ecology for its role in dispersing seeds and pollen.
The golden-bellied capuchin, also known as the yellow-breasted or buff-headed capuchin, is a species of New World or neotropical monkey. It lives mainly in trees and are omnivorous, eating a wide variety of both plant and animals as food. Golden-bellied capuchin normal home range is in the Atlantic forest of Brazil and it is critically endangered due to forest fragmentation and habitat loss mainly due to agriculture, there are currently efforts to protect them by the local government.
The capuchin monkeys are New World monkeys of the subfamily Cebinae. They are readily identified as the "organ grinder" monkey, and have been used in many movies and television shows. The range of capuchin monkeys includes some tropical forests in Central America and South America as far south as northern Argentina. In Central America, where they are called white-faced monkeys ("carablanca"), they usually occupy the wet lowland forests on the Caribbean coast of Costa Rica and Panama and deciduous dry forest on the Pacific coast.
White-fronted capuchin can refer to any of a number of species of gracile capuchin monkey which used to be considered as the single species Cebus albifrons. White-fronted capuchins are found in seven different countries in South America: Bolivia, Brazil, Colombia, Venezuela, Ecuador, Peru, and Trinidad and Tobago.
The Kaapori capuchin, also known as the Ka'apor capuchin, is a species of frugivorous, gracile capuchin endemic to the Brazilian Amazon. Their geographical home range is relatively small and is within the most densely populated region of the Amazon. With the strong human presence nearby,C. kaapori has had to deal with very large amounts of disturbances to their habitat and is one of the most endangered of the neotropical primates.
The wedge-capped capuchin or Guianan weeper capuchin is a capuchin monkey from South America. It is found in northern Brazil, Guyana and Venezuela. Cebus olivaceus is known to dwell in tall, primary forest and travel over long distances during the day.
The black capuchin, 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.
The black-striped capuchin, also known as the bearded capuchin, is a New World monkey in the family Cebidae. They can be found in northern and central Brazil. These capuchins mostly live in dry forests, and savannah landscapes between the Rio Araguaia and the Rio Grande. Known for its tool use, the black-striped capuchin has been shown to use tools in a wide variety of situations, ranging from using rocks for nut cracking to using sticks for digging. They were, until recently, considered a subspecies of the tufted capuchin, but because of more research and insights, they are considered their own species by many.
The large-headed capuchin is a subspecies of the tufted capuchin monkey from South America. It is found in Bolivia, Brazil, Colombia, Ecuador and Peru. It was formerly thought to be its own species, but studies have found it to be a subspecies of the tufted capuchin.
The blond capuchin is a species of capuchin monkey endemic to northeastern Brazil. This endangered species was rediscovered in 2006. It can live in exceptionally large groups of over 150 individuals, and like other capuchin species, exhibits a complex and high level of sociality. It is threatened by loss of habitat due to agriculture, primarily sugarcane fields. In many cases this has caused sugarcane to make up a large portion of their diet, which would otherwise consist of mostly fruit and small animals. The blond capuchin is known to inhabit both the Atlantic forest and Caatinga biomes, although the habitation of the Caatinga may be a recent choice caused by human encroachment into its former habitats. Like other primate species, the blond capuchin is also threatened by poaching and capture for the illegal pet trade.
Tool use by non-humans is a phenomenon in which a non-human animal uses any kind of tool in order to achieve a goal such as acquiring food and water, grooming, combat, defence, communication, recreation or construction. Originally thought to be a skill possessed only by humans, some tool use requires a sophisticated level of cognition. There is considerable discussion about the definition of what constitutes a tool and therefore which behaviours can be considered true examples of tool use. A wide range of animals, including mammals, birds, fish, cephalopods, and insects, are considered to use tools.
Primate cognition is the study of the intellectual and behavioral skills of non-human primates, particularly in the fields of psychology, behavioral biology, primatology, and anthropology.
Robust capuchin monkeys are capuchin monkeys in the genus Sapajus. Formerly, all capuchin monkeys were placed in the genus Cebus. Sapajus was erected in 2012 by Jessica Lynch Alfaro et al. to differentiate the robust (tufted) capuchin monkeys from the gracile capuchin monkeys, which remain in Cebus.
Gracile capuchin monkeys are capuchin monkeys in the genus Cebus. At one time all capuchin monkeys were included within the genus Cebus. In 2011, Jessica Lynch Alfaro et al. proposed splitting the genus between the robust capuchin monkeys, such as the tufted capuchin, and the gracile capuchins. The gracile capuchins retain the genus name Cebus, while the robust species have been transferred to Sapajus.
The crested capuchin or robust tufted capuchin is a species of robust capuchin monkey. It is endemic to Brazil. It was formerly considered a subspecies of the black capuchin but is now considered by some to be a separate species.
Azaras's capuchin or hooded capuchin is a species of robust capuchin. It occurs in eastern Paraguay, southeastern Bolivia, northern Argentina, and Brazil, at Mato Grosso do Sul and Mato Grosso states, in Pantanal. Its habitat consists of subtropical, humid, semi-deciduous, gallery forests and forested regions of the Pantanals. Formerly, it was considered a subspecies of black-striped capuchin, according to Groves (2005) with the name Cebus libidinosus paraguayanus, but Silva Jr. (2001) considered it a separated species. They are considered as frugivores-insectivores which means that their diet mainly consists of a variety of fruits, seeds, arthropods, frogs, small mammals, etc.
Social learning refers to learning that is facilitated by observation of, or interaction with, another animal or its products. Social learning has been observed in a variety of animal taxa, such as insects, fish, birds, reptiles, amphibians and mammals.
Primate archaeology is a field of research established in 2008 that combines research interests and foci from primatology and archaeology. The main aim of primate archaeology is to study behavior of extant and extinct primates and the associated material records. The discipline attempts to move beyond archaeology's anthropocentric perspective by placing the focus on both past and present primate tool use.
The Margarita Island capuchin, also known as the tufted Margarita Island capuchin or mono de margarita is a subspecies of the tufted capuchin that only resides in the Caribbean Sea on Margarita Island, Venezuela.
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