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{{Short description|Subfamily of New World monkeys}}
{{Automatic taxobox
 | name                = Capuchin monkey
 | image               = Capuchin Costa Rica.jpg
 | image_caption       = [[Panamanian white-faced capuchin]] (''Cebus imitator'') on a tree near a river bank in the jungles of [[Guanacaste Province|Guanacaste]], [[Costa Rica]]
 | fossil_range        = {{Fossil range|6.2|0}}Late [[Miocene]]-[[Holocene|Present]]
| taxon = Cebinae
| authority = [[Charles Lucien Bonaparte|Bonaparte]], 1831
| subdivision_ranks = Genera
| subdivision = 
''[[gracile capuchin monkey|Cebus]]''<br />
''[[robust capuchin monkey|Sapajus]]''
}}

The '''capuchin monkeys''' ({{IPAc-en|ˈ|k|æ|p|j|ʊ|(|t|)|ʃ|ɪ|n}}) are [[New World monkey]]s of the subfamily '''Cebinae'''. They are readily identified as the "[[Street organ|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 [[Geography of Costa Rica|Caribbean coast of Costa Rica]] [[Geography of Panama|and Panama]] and deciduous dry forest on the Pacific coast.

== Etymology ==

The word "capuchin" derives from the [[Order of Friars Minor Capuchin]], who wear brown robes with large hoods. When [[Portuguese maritime exploration|Portuguese explorers]] reached the [[Americas]] in the 15th century, they found small monkeys whose coloring resembled these friars, especially when in their robes with hoods down, and named them capuchins.<ref name=Fragaszy/> When the scientists described a specimen (thought to be a [[golden-bellied capuchin]]) they noted that: "his muzzle of a tanned color, ... with the lighter color around his eyes that melts into the white at the front, his cheeks ..., give him the looks that involuntarily reminds us of the appearance that historically in our country represents ignorance, laziness, and sensuality."<ref>{{cite book |author1=Saint-Hilaire, E. G.  |author2=Cuvier, F. G. |year=1924 |title=Histoire Naturelle des Mammifères |oclc=166026273 |publisher=Paris, impr. de C. de Lasteyrie}}</ref> The scientific name of the genus, ''Cebus'' comes from the Greek word'' kêbos'',<ref>{{cite book |author=Rossiter, William |year=1879 |title=An illustrated dictionary of scientific terms |publisher=William Collins, Sons, and Company |location=London & Glasgow |isbn=978-0-548-93307-7}}</ref> meaning a long-tailed monkey.

== Classification ==

The species-level [[Taxonomy (biology)|taxonomy]] of this subfamily remains highly controversial, and alternative treatments than the one listed below have been suggested.<ref>{{cite journal | last1 = Amaral | first1 = P. J. S | last2 = Finotelo | first2 = L. F. M. | last3 = De Oliveira | first3 = E. H. C | last4 = Pissinatti | first4 = A. | last5 = Nagamachi | first5 = C. Y. | last6 = Pieczarka | first6 = J. C. | year = 2008 | title = Phylogenetic studies of the genus ''Cebus'' (Cebidae-Primates) using chromosome painting and G-banding. | journal = BMC Evol. Biol. | volume = 8 | issue = 1 | page = 169 | doi=10.1186/1471-2148-8-169 | pmid=18534011 | pmc=2435554 | doi-access = free | bibcode = 2008BMCEE...8..169A }}</ref><ref>{{cite journal | last1 = Rylands | first1 = A. B. | last2 = Kierulff | first2 = M. C. M. | last3 = Mittermeier | first3 = R. A. | year = 2005 | title = Notes on the taxonomy and distributions of the tufted capuchin monkeys (''Cebus'', Cebidae) of South America | url =https://www2.icb.ufmg.br/lundiana/Contents/full/vol6sup2005/17.pdf | journal = Lundiana | volume = 6 | issue = supp | pages = 97–110 | doi = 10.35699/2675-5327.2005.22130 }}</ref><ref name=silva2001>Silva Jr., J. de S. (2001). ''Especiação nos macacos-prego e caiararas, gênero ''Cebus'' Erxleben, 1777 (Primates, Cebidae).'' PhD thesis, Rio de Janeiro, Universidade Federal do Rio de Janeiro.</ref><ref name=iucn>[[International Union for Conservation of Nature|IUCN]] (2008). [http://www.iucnredlist.org/ 2008 IUCN Red List of Threatened Species.] Accessed 23 November 2008</ref>

In 2011, Jessica Lynch Alfaro et al. proposed that the [[robust capuchin]]s (formerly the ''C. apella'' group) be placed in a separate genus, ''Sapajus'', from the [[gracile capuchin]]s (formerly the ''C. capucinus'' group) which retain the genus ''Cebus''.<ref name="alfaro-genetic">{{cite journal|title=Explosive Pleistocene range expansion leads to widespread Amazonian sympatry between robust and gracile capuchin monkeys|author=Lynch Alfaro, J.W.|journal=Journal of Biogeography|url=http://xa.yimg.com/kq/groups/19113390/1559919033/name/jbi_2609_Rev_EVcorrectedproof.pdf|doi=10.1111/j.1365-2699.2011.02609.x|year=2011|display-authors=etal|volume=39|issue=2|pages=272–288|s2cid=13791283 |url-status=dead|archive-url=https://web.archive.org/web/20150226082115/http://xa.yimg.com/kq/groups/19113390/1559919033/name/jbi_2609_Rev_EVcorrectedproof.pdf|archive-date=2015-02-26}}</ref><ref name=alfaro>{{cite journal|title=How Different Are Robust and Gracile Capuchin Monkeys? An Argument for the Use of ''Sapajus'' and ''Cebus''|author1=Lynch Alfaro, J.W. |author2=Silva, j. |author3=Rylands, A.B. |doi=10.1002/ajp.22007|pmid=22328205|year=2012|pages=1–14|journal=American Journal of Primatology |volume=74|issue=4 |s2cid=18840598 }}</ref> Other primatologists, such as [[Paul Garber]], have begun using this classification.<ref>{{cite journal|title=Experimental Field Study of Problem-Solving Using Tools in Free-Ranging Capuchins (''Sapajus nigritus'', formerly ''Cebus nigritus'')|author=Garber, P.A.|author-link=Paul Garber|author2=Gomes, D.F.|author3=Bicca-Marquez, J.C.|name-list-style=amp|journal=American Journal of Primatology|volume=74|year=2011|pages=344–58|pmid=21538454|url=https://netfiles.uiuc.edu/p-garber/www/publications/Garberetal2011Capuchintool.pdf|archive-url=https://web.archive.org/web/20121218211740/https://netfiles.uiuc.edu/p-garber/www/publications/Garberetal2011Capuchintool.pdf|url-status=dead|archive-date=2012-12-18|doi=10.1002/ajp.20957|issue=4|s2cid=39363765}}</ref>

According to genetic studies led by Lynch Alfaro in 2011, the gracile and robust capuchins diverged approximately 6.2 million years ago. Lynch Alfaro suspects that the divergence was triggered by the creation of the Amazon River, which separated the monkeys in the Amazon north of the Amazon River, who then evolved into the gracile capuchins. Those in the Atlantic Forest south of the river evolved into the robust capuchins. Gracile capuchins have longer limbs relative to their body size than robust capuchins, and have rounder skulls, whereas robust capuchins have jaws better adapted for opening hard nuts. Robust capuchins have crests and the males have beards.<ref name="alfaro-genetic" /><ref name="alfaro" />

* '''Genus ''[[Cebus]]'''''<ref name=MSW3>{{MSW3 Groves|pages=136–138|id=12100255}}</ref><ref name=handbook>{{cite book|title=Handbook of the Mammals of the World: Volume 3, Primates|year=2012|isbn=978-8496553897|publisher=Lynx |editor=Mittermeier, Russell A. |editor2=Rylands, Anthony B. |editor3=Wilson, Don E.|pages=412–413}}</ref>
** [[Colombian white-faced capuchin]] or Colombian white-headed capuchin, ''Cebus capucinus''
** [[Panamanian white-faced capuchin]] or Panamanian white-headed capuchin, ''Cebus imitator''
** [[Marañón white-fronted capuchin]], ''Cebus yuracus''
** [[Shock-headed capuchin]], ''Cebus cuscinus''
** [[Spix's white-fronted capuchin]], ''Cebus unicolor''
** [[Humboldt's white-fronted capuchin]], ''Cebus albifrons''
** [[Guianan weeper capuchin]], ''Cebus olivaceus''
**[[Chestnut weeper capuchin]], ''Cebus castaneus''
** [[Ka'apor capuchin]], ''Cebus kaapori''
** [[Venezuelan brown capuchin]], ''Cebus brunneus''
** [[Sierra de Perijá white-fronted capuchin]], ''Cebus leucocephalus''
** [[Río Cesar white-fronted capuchin]], ''Cebus cesare''
** [[Varied white-fronted capuchin]], ''Cebus versicolor''
** [[Santa Marta white-fronted capuchin]], ''Cebus malitiosus''
** [[Ecuadorian white-fronted capuchin]], ''Cebus aequatorialis''

[[File:Capuchin monkeys sharing.jpg|thumb|upright|[[Tufted capuchin]] (''Sapajus apella'')]]

* '''Genus ''[[Sapajus]]'''''<ref name=silva2001/><ref name=alfaro/>
** Black-capped, brown or [[tufted capuchin]], ''Sapajus apella''
*** [[Guiana brown capuchin]], ''Sapajus apella apella''
*** ''Sapajus apella fatuellus''
*** [[Large-headed capuchin]], ''Sapajus apella macrocephalus''
*** Margarita Island capuchin, ''Sapajus apella margaritae''
*** ''Sapajus apella peruanus''
*** ''Sapajus apella tocantinus''
** [[Blond capuchin]], ''Sapajus flavius''*
** [[Black-striped capuchin]], ''Sapajus libidinosus''
*** ''Sapajus libidinosus juruanus''
*** ''Sapajus libidinosus libidinosus''
*** ''Sapajus libidinosus pallidus''
*** ''Sapajus libidinosus paraguayanus''
** [[Azaras's capuchin]], ''Sapajus cay''
**[[Black capuchin]], ''Sapajus nigritus''
*** ''Sapajus nigritus cucullatus''
*** ''Sapajus nigritus nigritus''
** [[Crested capuchin]] or robust tufted capuchin, ''Sapajus robustus''
** [[Golden-bellied capuchin]], ''Sapajus xanthosternos''

<nowiki>*</nowiki> Rediscovered species.<ref>{{cite journal | journal = Boletim do Museu Nacional (Rio de Janeiro), Zoologia |series=Nova Série | url = http://acd.ufrj.br/~museuhp/CP/Bol-Zool/BolZool2006/Bol%20Zool%20MN%20523.pdf | issue = 523 | year = 2006 | pages = 1–16 | title = Rediscovery of Marcgrave's capuchin monkey and designation of a neotype for ''Simia flavia'' Schreber, 1774 (Primates, Cebidae) | author1 = de Oliveira, M. M. | author2 = Langguth, A. | url-status = dead | archive-url = https://web.archive.org/web/20090205133225/http://acd.ufrj.br/~museuhp/CP/Bol-Zool/BolZool2006/Bol%20Zool%20MN%20523.pdf | archive-date = 2009-02-05 }} See also: {{cite journal | journal = Zootaxa | url = http://www.mapress.com/zootaxa/2006f/zt01200p012.pdf | issue = 1200 | year = 2006 | pages = 1–12 | title = A new species of capuchin monkey, genus ''Cebus'' Erxleben (Cebidae, Primates): found at the very brink of extinction in the Pernambuco Endemism Centre | author = Mendes Pontes, A. R. | author2 = Malta, A. | author3 = Asfora, P. H. | name-list-style = amp }}</ref>

The oldest known crown [[Platyrrhini|platyrrhine]] and member of Cebidae, ''[[Panamacebus transitus]]'', is estimated to have lived 21 million years ago. It is the earliest known fossil evidence of a mammal travelling between South and North America.<ref>{{cite journal |title=First North American fossil monkey and early Miocene tropical biotic interchange |year=2016 |last1=Bloch |first1=Jonathan I. |last2=Woodruff |first2=Emily D. |last3=Wood |first3=Aaron R. |last4=Rincon |first4=Aldo F. |last5=Harrington |first5=Arianna R. |last6=Morgan |first6=Gary S. |last7=Foster |first7=David A. |last8=Montes |first8=Camilo |last9=Jaramillo |first9=Carlos A. |last10=Jud |first10=Nathan A. |last11=Jones |first11=Douglas S. |last12=MacFadden |first12=Bruce J. |journal=Nature |volume=533 |issue=7602 |pages=243–246 |pmid=27096364 |s2cid=4445687 |doi=10.1038/nature17415|bibcode=2016Natur.533..243B }}</ref>

== Physical characteristics ==

Capuchins are black, brown, [[buff (color)|buff]] or whitish, but their exact color and pattern depends on the species involved. Capuchin monkeys are usually dark brown with a cream/off-white coloring around their necks. They reach a length of {{convert|30|to|56|cm|in|round=0.5|abbr=on}}, with tails that are just as long as the body. On average, they weigh from 1.4 to 4&nbsp;kg (3 to 9 pounds) and live up to 25 years old in their natural habitats,{{citation needed|date=September 2022}} and up to 35 in captivity.{{citation needed|date=September 2022}}

== Habitat and distribution ==

Capuchins prefer environments that give them access to shelter and easy food, such as low-lying forests, mountain forests, and rain forests. They are particularly abundant in Argentina, Brazil, Costa Rica, Honduras, Paraguay, and Peru. They use these areas for shelter at night and food access during the day. The canopy of the trees allows for protection from threats above, and the capuchin monkeys' innate ability to climb trees with ease allows them to escape and hide from predators on the jungle floor. This environment is mutually beneficial for the capuchins and for the ecosystem in which they inhabit. This is because they spread their seed leftovers and fecal matter across the [[forest floor]] which helps new plants to grow, therefore adding to the already abundant foliage that shelters the capuchin.{{citation needed|date=November 2019}}

== Behavior ==
[[File:Capuchin Monkey enjoys a massage.jpg|thumb|A capuchin monkey (left) enjoying a massage from another capuchin monkey.]]
Like most New World monkeys, capuchins are [[diurnality|diurnal]] and [[arboreal locomotion|arboreal]]. Capuchins are polygamous, and the females mate throughout the year, but only go through a gestation period once every 2 years between December and April. Females bear young every two years following a 160- to 180-day gestation.{{citation needed|date=November 2019}} The young cling to their mother's chest until they are larger, then they move to her back. Adult male capuchin rarely take part in caring for the young. Juveniles become fully mature within four years for females and eight years for males. In captivity, individuals have reached an age of 50 years, although natural life expectancy is only 15 to 25 years. Capuchins live in groups of 6–40 members, consisting of related females, their offspring, and several males.{{citation needed|date=November 2019}}

[[File:Cebus albifrons edit.jpg|thumb|upright|left|White-fronted capuchin (''Cebus albifrons'')]]

=== Diet ===

The capuchin monkey feeds on a vast range of food types, and is more varied than other monkeys in the family [[Cebidae]]. They are [[omnivores]], and consume a variety of plant parts such as leaves, flower and fruit, seeds, pith, woody tissue, sugarcane, bulb, and exudates, as well as [[arthropods]], [[molluscs]], a variety of [[vertebrates]], and even [[primates]].<ref name="Izawa K 1979">{{cite journal | last1 = Izawa | first1 = K | year = 1979 | title = Foods and feeding behaviour of wild black-capped capuchin (''Cebus apella'') | journal = Primates | volume = 20 | pages = 57–76 | doi=10.1007/bf02373828| s2cid = 30424050 }}</ref> Recent findings of old stone tools in Capuchin habitats have suggested that recently the Capuchins have switched from small nuts, such as cashews, to larger and harder nuts. Capuchins have also been observed to be particularly good at catching frogs.<ref name="Izawa K 1979"/> They are characterized as innovative and extreme foragers because of their ability to acquire sustenance from a wide collection of unlikely food, which may assure their survival in habitats with extreme food limitation.<ref>{{cite book|author1=Fragaszy, Dorothy M. |author2=Visalberghi, Elisabetta |author3=Fedigan, Linda M. |title=The Complete Capuchin: The Biology of the Genus ''Cebus''|chapter-url=https://books.google.com/books?id=A6TmtS_qOwgC&pg=PA36|date=21 June 2004|publisher=Cambridge University Press|isbn=978-0-521-66768-5|pages=36–|chapter=Behavioral ecology: how do capuchins make a living?}}</ref> Capuchins living near water will also eat crabs and shellfish by cracking their shells with stones.<ref>{{cite journal | last1 = Port-Carvalhoa | first1 = M. | last2 = Ferraria | first2 = S. F. | last3 = Magalhãesc | first3 = C. | year = 2004 | title = Predation of Crabs by Tufted Capuchins (''Cebus apella'') in Eastern Amazonia | journal = Folia Primatol | volume = 75 | issue = 3| pages = 154–156 | doi=10.1159/000078305 | pmid=15240980| s2cid = 1647323 }}</ref>

=== Social structure ===

Capuchin monkeys often live in large groups of 10 to 35 individuals within the forest, although they can easily adapt to places colonized by humans. The Capuchins have discrete hierarchies that are distinguished by age and sex.<ref>{{Cite web|url=http://pin.primate.wisc.edu/factsheets/entry/tufted_capuchin/behav|title=Primate Factsheets: Tufted capuchin (''Cebus apella'') Behavior|website=pin.primate.wisc.edu|access-date=2019-11-12}}</ref> Usually, a single male will dominate the group, and he will have primary rights to [[Mating|mate]] with the females of the group. However, the [[white-headed capuchin]] groups are led by both an alpha male and an alpha female.<ref>{{cite book|vauthors=Ferrari SF, Iwanaga S, Ravetta AL, Freitas FC, Sousa BA, Souza LL, Costa CG, Coutinho PE |year=2003|chapter= Dynamics of Primate Communities along the Santarém-Cuiabá Highway in South-Central Brazilian Amazonia|editor=Marsh LK |title=Primates in Fragments|publisher= Kluwer|place= New York|pages=123–144|doi=10.1007/978-1-4757-3770-7_9|isbn=978-1-4757-3770-7}}</ref> Each group will cover a large territory, since members must search for the best areas to feed. These primates are territorial animals, distinctly marking a central area of their territory with urine and defending it against intruders, though outer areas may overlap. The stabilization of group dynamics is served through mutual grooming, and communication occurs between the monkeys through various calls.<ref>{{cite journal|doi=10.1007/s00265-012-1421-5|title=Collective group movement and leadership in wild black howler monkeys (''Alouatta pigra'')|journal=Behavioral Ecology and Sociobiology|volume=67|pages=31–41|year=2012|last1=Van Belle|first1=Sarie|last2=Estrada|first2=Alejandro|last3=Garber|first3=Paul A.|s2cid=14844073}}
</ref> Their vocal communications have various meanings such as creating contact with one another, warning about a predator, and forming new groups. The social experience of the capuchins directly influences the development of attention in society. They create new social behaviors within multiple groups that signify different types of interactions. These include; tests of friendship, displays against enemies, infant and sexual intimacy.<ref>{{Cite web|url=https://www.sciencedaily.com/releases/2017/08/170824094042.htm|title=How new behaviors appear and spread among capuchin monkeys|website=ScienceDaily|language=en|access-date=2019-11-12}}</ref> This creates social rituals that are designed to test the strength of social bonds and a reliance on social learning.<ref>{{Cite journal|last1=Fragaszy|first1=Dorothy M.|last2=Eshchar|first2=Yonat|last3=Visalberghi|first3=Elisabetta|last4=Resende|first4=Briseida|last5=Laity|first5=Kellie|last6=Izar|first6=Patrícia|date=2017-07-25|title=Synchronized practice helps bearded capuchin monkeys learn to extend attention while learning a tradition|journal=Proceedings of the National Academy of Sciences|language=en|volume=114|issue=30|pages=7798–7805|doi=10.1073/pnas.1621071114|issn=0027-8424|pmid=28739944|pmc=5544277|bibcode=2017PNAS..114.7798F |doi-access=free}}</ref>

=== Mating ===

Capuchin females often direct most of their proceptive and mating behavior towards the alpha male. However, when the female reaches the end of her proceptive period, she may sometimes mate with up to six different subordinate males in one day.<ref>{{cite journal | last1 = Janson | first1 = C. H. | year = 1984 | title = Female choice and mating system of the brown capuchin monkey ''Cebus apella'' (Primates: Cebidae) | journal = Zeitschrift für Tierpsychologie | volume = 65 | issue = 3| pages = 177–200 | doi=10.1111/j.1439-0310.1984.tb00098.x| bibcode = 1984Ethol..65..177J }}</ref> Strictly targeting the alpha male does not happen every time, as some females have been observed to mate with three to four different males.<ref>{{cite journal | last1 = Lynch | first1 = J. W. | year = 1998 | title = Mating behavior in wild tufted capuchins (''Cebus apella nigritus'') in Brazil's Atlantic forest | journal = Am. J. Phys. Anthropol. | volume = 105 | issue = Suppl. 26 | doi = 10.1002/(SICI)1096-8644(1998)26+<148::AID-AJPA13>3.0.CO;2-U | page = 153 }}</ref> When an alpha female and a lower-ranking female want to mate with an alpha male, the more dominant female will get rights to the male over the lower-ranking one.

== Intelligence ==
[[File:Cebus robustus (Crested capuchin monkey).jpg|thumb|right|[[Crested capuchin]] (''Sapajus robustus'')]]

The capuchin is considered to be the most intelligent New World monkey<ref>{{cite web |url=http://www.monkeyjungle.com/amazonian.htm |title=Black-faced Capuchin |work=Amazonian Rainforest |publisher=Monkey Jungle |access-date=2008-10-13}}</ref> and is often kept in captivity. The tufted monkey is especially noted for its long-term tool usage,<ref>{{cite web|title=Saving the monkeys|url=https://spie.org/membership/spie-professional-magazine/journal-of-applied-remote-sensing-saving-monkey-habitat|publisher=SPIE Professional|access-date=1 January 2016}}</ref> one of the few examples of primate tool use other than by [[ape]]s including humans. Upon seeing [[macaw]]s eating [[Oil palm|palm nuts]], cracking them open with their beaks, this monkey will select a few of the ripest fruits, nip off the tip of the fruit and drink down the juice, then seemingly discard the rest of the fruit with the nut inside. When these discarded fruits have hardened and become slightly brittle, the capuchin will gather them up again and take them to a large flat boulder where they have previously gathered a few river stones from up to a mile away. They will then use these stones, some of them weighing as much as the monkeys, to crack open the fruit to get to the nut inside. Young capuchins will watch this process to learn from the older, more experienced adults but it takes them 8 years to master this.<ref>{{cite journal |title=Substrate and Tool Use by Brown Capuchins in Suriname: Ecological Contexts and Cognitive Bases |author=Boinski, S. |author2=Quatrone, R. P. |author3=Swartz, H. |name-list-style=amp |year=2008 |volume=102 |issue=4 |pages=741–761 |doi=10.1525/aa.2000.102.4.741 |journal=American Anthropologist}}</ref> The learning behavior of capuchins has been demonstrated to be directly linked to a reward rather than curiosity.<ref>{{Cite journal|last1=Edwards|first1=Brian J.|last2=Rottman|first2=Benjamin M.|last3=Shankar|first3=Maya|last4=Betzler|first4=Riana|last5=Chituc|first5=Vladimir|last6=Rodriguez|first6=Ricardo|last7=Silva|first7=Liara|last8=Wibecan|first8=Leah|last9=Widness|first9=Jane|date=2014-02-19|title=Do Capuchin Monkeys (''Cebus apella'') Diagnose Causal Relations in the Absence of a Direct Reward?|journal=PLOS ONE|volume=9|issue=2|pages=e88595|doi=10.1371/journal.pone.0088595|issn=1932-6203|pmc=3929502|pmid=24586347|bibcode=2014PLoSO...988595E|doi-access=free}}</ref>

In 2005, experiments were conducted on the ability of capuchins to use money.<ref name="anderson.ucla.edu">{{cite journal|last1=Chen|first1=M. Keith|last2=Lakshminarayanan|first2=Venkat|last3=Santos|first3=Laurie R.|title=How Basic Are Behavioral Biases? Evidence from Capuchin Monkey Trading Behavior|journal=Journal of Political Economy|date=2006|volume=114|issue=3|pages=517–537|doi=10.1086/503550|url=http://www.anderson.ucla.edu/faculty/keith.chen/papers/Final_JPE06.pdf|citeseerx=10.1.1.594.4936|s2cid=18753437}}</ref> After several months of training, the monkeys began exhibiting behaviors considered to reflect an understanding of the concept of a medium of exchange that were previously believed to be restricted to humans (such as responding rationally to price shocks).<ref name="anderson.ucla.edu"/> They showed the same propensity to [[loss aversion|avoid perceived losses]] demonstrated by human subjects and investors.

During the mosquito season, they crush [[millipede]]s and rub the result on their backs. This acts as a natural [[insect repellent]].<ref>{{Cite journal |title=Seasonal Anointment with Millipedes in a Wild Primate: A Chemical Defense Against Insects? |journal=Journal of Chemical Ecology |volume=26 |issue=12 |year=2000 |doi=10.1023/A:1026489826714 |pages=2781–2790 |author=Valderrama, X.|bibcode=2000JCEco..26.2781V |s2cid=25147071 |display-authors=etal}}</ref>

=== Self-awareness ===
{{Further|Self-awareness}}

When presented with a reflection, capuchin monkeys react in a way that indicates an intermediate state between seeing the mirror as another individual and recognizing the image as self.

Most animals react to seeing their reflections as if encountering another individual they do not recognize. An experiment with capuchins shows that they react to a reflection as a strange phenomenon, but not as if seeing a strange capuchin.

In the experiment, capuchins were presented with three different scenarios:
# Seeing an unfamiliar, same-sex monkey on the other side of a clear barrier.
# Seeing a familiar, same-sex monkey on the other side of a clear barrier.
# A mirror showing a reflection of the monkey.

In scenario 1, females appeared anxious and avoided eye-contact, while males made threatening gestures. In scenario 2, there was little reaction by either males or females.

When presented with a reflection, females gazed into their own eyes and made friendly gestures, such as lip-smacking and swaying. Males made more eye contact than with strangers or familiar monkeys but reacted with signs of confusion or distress, such as squealing, curling up on the floor, or trying to escape from the test room.<ref>{{cite journal | author=de Waal, F. B. | author2=Dindo, M. | author3=Freeman, C. A. | author4=Hall, M. J. | name-list-style=amp | title=The monkey in the mirror: Hardly a stranger | journal=[[Proceedings of the National Academy of Sciences]] | volume=102 | issue= 32| year=2005 | pages= 11140–7| pmid=16055557 | doi=10.1073/pnas.0503935102 | pmc=1183568| bibcode=2005PNAS..10211140D | doi-access=free }}</ref>

=== Theory of mind ===
{{Main|Theory of mind}}

The question of whether capuchin monkeys have a theory of mind—whether they can understand what another creature may know or think—has been neither proven nor disproven conclusively. If confronted with a ''knower-guesser'' scenario, where one trainer can be observed to know the location of food and another trainer merely guesses the location of food, capuchin monkeys can learn to rely on the knower.<ref>{{cite journal | last = Kuroshima | first = Hika |author2=Fujita, Kazuo |author3=Fuyuki, Akira |author4=Masuda, Tsuyuka |date=March 2002 | title = Understanding of the relationship between seeing and knowing by tufted capuchin monkeys (''Cebus apella'') | journal = Animal Cognition | volume = 5 | issue = 1 | pages = 41–48 | doi = 10.1007/s10071-001-0123-6  | pmid = 11957401 | s2cid = 10783449 | issn = 1435-9448 }}</ref> This has, however, been repudiated as conclusive evidence for a theory of mind as the monkeys may have learned to discriminate knower and guess by other means.<ref>{{cite journal|author=Heyes, C. M. |year=1998 |title=Theory Of Mind In Nonhuman Primates |journal=Behavioral and Brain Sciences|doi=10.1017/S0140525X98000703 |volume=21 |issue=1 |pmid=10097012 |pages=101–14 |s2cid=6469633 |url=http://discovery.ucl.ac.uk/117063/1/download.14pdf.pdf }}</ref> Until recently it was believed that non-human [[hominidae|great ape]]s did not possess a theory of mind either, although recent research indicates this may not be correct.<ref>{{cite web|last=Jabr|first=Ferris|title=Clever critters: Bonobos that share, brainy bugs and social dogs|url=http://blogs.scientificamerican.com/observations/2010/06/08/clever-critters-bonobos-that-share-brainy-bugs-and-social-dogs/|publisher=Scientific American|date=8 June 2010}}</ref> Human children commonly develop a theory of mind around the ages 3 and 4.

== Threats ==

Capuchin monkeys are threatened by deforestation, the [[Pet monkey|pet trade]], and humans hunting for bushmeat.<ref name=iucn/> According to the [[IUCN Red List|IUCN Red List of Threatened Species]], nearly all species are decreasing in population, with many facing threats of extinction.<ref>{{Cite web |date=August 26, 2024 |title=Search for "Capuchin" |url=https://www.iucnredlist.org/search?query=Capuchin&searchType=species |access-date=August 26, 2024 |website=IUCN Red List Threatened Species}}</ref> Since capuchins have a high reproductive rate and can adapt to different living environments, they can survive forest loss more than some other species; however, [[habitat fragmentation]] is still a threat.<ref>{{cite journal | last1 = Sanz | first1 = V | last2 = Márquez | first2 = L | year = 1994 | title = Conservación del mono capuchino de Margarita (''Cebus apella margaritae'') en la Isla de Margarita, Venezuela | url = https://www.researchgate.net/publication/263965664| journal = Neotropical Primates | volume = 2 | issue = 2| pages = 5–8 | doi = 10.62015/np.1994.v2.203 | doi-access = free }}</ref> Predators include jaguars, cougars, [[jaguarundi]]s, coyotes, [[tayra]]s, snakes, crocodiles, birds of prey, and humans. The main predator of the [[tufted capuchin]] is the [[harpy eagle]], which has been seen bringing several capuchin back to its nest.<ref name=Fragaszy>{{cite book|author1=Fragaszy, Dorothy M. |author2=Visalberghi, Elisabetta |author3=Fedigan, Linda M. |title=The Complete Capuchin: The Biology of the Genus ''Cebus''|url=https://books.google.com/books?id=A6TmtS_qOwgC|date=21 June 2004|publisher=Cambridge University Press|isbn=978-0-521-66768-5|page=5}}</ref>

== Relationship with humans ==

[[File:Organ grinder with monkey.jpg|upright|thumb|200px|19th-century organ grinder and his capuchin monkey]]
Easily recognized as the "[[organ grinder]]" or "[[greyhound jockey]]" monkeys, capuchins are sometimes kept as [[Exotic pet#Primates|exotic pets]]. Sometimes they plunder fields and crops and are seen as troublesome by nearby human populations.<ref name="Fragaszy" /> In some regions, they have become rare due to the destruction of their habitat.<ref name="Fragaszy" />

Capuchins have been used as service animals, and were once referred to as "nature's butlers" by the AARP.<ref name = AARP>{{cite web |url=http://www.aarp.org/makeadifference/volunteer/articles/monkey_do.html|archive-url=https://web.archive.org/web/20080916115914/http://www.aarp.org/makeadifference/volunteer/articles/monkey_do.html|url-status=dead|archive-date=2008-09-16|title=Animals in Service|author=Lineberry, Cate |publisher=[[AARP]] |access-date=2008-10-13 }}</ref> Helping Hands, a nonprofit organization, trained capuchin monkeys to assist [[Quadriplegia|quadriplegics]] as [[Service animal#Helper monkey|monkey helper]]s in a manner similar to [[mobility assistance dog]]s.<ref name=AARP/>

In 2010, the U.S. federal government revised its definition of service animal under the Americans with Disabilities Act (ADA). Non-human primates are no longer recognized as service animals under the ADA.<ref name="ada">{{cite web|title=Highlights of the Final Rule to Amend the Department of Justice's Regulation Implementing Title II of the ADA|url=http://www.ada.gov/regs2010/factsheets/title2_factsheet.html|publisher=United States Department of Justice-Civil Rights Division|access-date=October 2, 2013|archive-url=https://web.archive.org/web/20180721221807/https://www.ada.gov/regs2010/factsheets/title2_factsheet.html|archive-date=July 21, 2018|url-status=dead}}</ref> The American Veterinary Medical Association does not support the use of nonhuman primates as assistance animals because of animal welfare concerns, the potential for serious injury to people, and risks that primates may transfer dangerous diseases to humans.<ref name="avma">{{cite web|title=AVMA Animal Welfare Division Director's Testimony on the Captive Primate Safety Act|url=https://www.avma.org/News/PressRoom/Pages/Testimony-golab-080311.aspx|publisher=American Veterinary Medicine Association|access-date=October 2, 2013|archive-url=https://web.archive.org/web/20180721192412/https://www.avma.org/News/PressRoom/Pages/Testimony-golab-080311.aspx|archive-date=July 21, 2018|url-status=dead}}</ref> In 2021, Helping Hands (the organization that provided helper monkeys to disabled persons) rebranded, changing its name to Envisioning Access and replaced the use of monkeys with a focus on new assistive technologies.<ref>{{Cite web |title=Envisioning Access |url=https://www.envisioningaccess.org/monkey-helpers-our-past/our-history/ |access-date=2024-06-18 |website=Envisioning Access |language=en-US}}</ref>

Capuchin monkeys are the most common featured [[monkeys]] in film and television, with notable examples including: ''[[Night at the Museum]]'' ([[Night at the Museum (film series)|and its sequels]]), ''[[Outbreak (1995 film)|Outbreak]]'', ''[[Monkey Shines (film)|Monkey Shines]]'', ''[[Pirates of the Caribbean: The Curse of the Black Pearl]]'' ([[Pirates of the Caribbean (film series)|and its sequels]]), ''[[Zookeeper (film)|Zookeeper]]'', ''[[George of the Jungle (film)|George of the Jungle]]'', and ''[[The Hangover Part II]]''. [[Ross Geller]] ([[David Schwimmer]]) on the [[NBC]] sitcom ''[[Friends]]'' had a capuchin monkey named Marcel. [[Crystal the Monkey]] is a famous monkey actress.

== References ==
{{Reflist|2}}

== External links ==
{{EB1911 poster|Capuchin Monkey}}
{{Wikispecies-inline|Cebinae}}<br/>
{{Commons category-inline|Cebinae}}
* [http://pin.primate.wisc.edu/factsheets/links/cebus Primate Info Net ''Cebus'' Factsheets]

{{Cebidae nav}}
{{Taxonbar|from=Q326260}}

[[Category:Capuchin monkeys| ]]
[[Category:Tool-using mammals]]
[[Category:Extant Miocene first appearances]]
<!-- [[Category:Animal subfamilies]] moved to Cebinae redirect-->