{{Short description|Species of primate}} {{Speciesbox | name = Panamanian white-faced capuchin | image = Panamanian white-faced capuchins (81622)2.jpg | status = VU | status_system = IUCN3.1 | status_ref = <ref name=iucn>{{cite iucn |author=Williams-Guillén, K. |author2=Rosales-Meda, M. |author3=Méndez-Carvajal, P.G.|author4=Solano-Rojas, D. |author5=Urbani, B |author6=Lynch-Alfaro, J.W. |date=2021 |title=''Cebus imitator'' |volume=2021 |article-number=e.T81265980A191708420 |doi=10.2305/IUCN.UK.2021-1.RLTS.T81265980A191708420.en |access-date=29 November 2021}}</ref> | genus = Cebus | species = imitator | authority = (Thomas, 1903) | range_map = Cebus capucinus and imitator Range Map.png | range_map_caption = Distribution of ''Cebus imitator'' and ''Cebus capucinus''.<ref name=tax/> ''Cebus imitator'' covers the Central American portion of the range except the easternmost portion of Panama. }}
The '''Panamanian white-faced capuchin''' ('''''Cebus imitator'''''), 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 Panamanian white-faced capuchin is versatile, living in many different types of forest, and eating many different types of food, including fruit, other plant material, invertebrates, and small vertebrates. It lives in troops that can exceed 20 animals and include both males and females. It is noted for its tool use, including rubbing plants over its body in an apparent use of herbal medicine, and also using tools as weapons and for getting to food. It is a long-lived monkey, with a maximum recorded age of over 54 years.
The white-faced capuchin is a medium-sized monkey, weighing up to {{convert|3.9|kg|lboz|abbr=on}}. It is mostly black, with a pink face and white on much of the front part of the body, giving it its common name. It has a distinctive prehensile tail that is often carried coiled up and is used to help support the monkey when it is feeding beneath a branch.
Panamanian white-faced capuchins are highly social, living in groups of 16 individuals on average, about three quarters of which are females. Groups consists of related females, immigrant males, and offspring. On average, females birth offspring every 27 months even though they mate throughout the year. Females tend to stay within their original group while males leave their natal group when they are four years old and change groups every four years thereafter. Both male and female capuchins exhibit different dominance behaviors within the group.
In captivity, the Panamanian white-faced capuchin has been used as a companion to the organ grinder. It is an intelligent monkey and has been portrayed as a pet by American media (for example, in the ''Pirates of the Caribbean'' film series).
== Taxonomy == [[File:White-faced Capuchin 1.jpeg|thumb|The Panamanian white-faced capuchin was previously considered a subspecies of the Colombian white-headed capuchin, ''Cebus capucinus imitator''.{{clarify|date=January 2019|reason=which species is pictured?}}]] [[File:Panamanian white-faced capuchin (Cebus imitator).jpg|thumb|in Gatun Lake, Panama]] The Panamanian white-faced capuchin is a member of the family Cebidae, the family of New World monkeys containing capuchin monkeys and squirrel monkeys. Until the 21st century, the Panamanian white-faced capuchin was considered conspecific with ''Cebus capucinus'', the Colombian white-faced capuchin, but as a separate subspecies, ''C. capucinus imitator''.<ref name=groves2>{{MSW3 Groves|page=Cebus|id=12100255}}</ref> Some primatologists continue to consider the Panamanian and Colombian white-faced capuchins as a single species.<ref name=handbook>{{cite book|title=Handbook of the Mammals of the World: Volume 3, Primates|year=2013|isbn=978-84-96553-89-7|publisher=Lynx|editor1= Mittermeier, Russell A. |editor2=Rylands, Anthony B. |editor3=Wilson, Don E. |author=Mittermeier, Russell A. |author2=Rylands, Anthony B. |pages=412–413}}</ref> The Panamanian monkey is a member of the ''C. capucinus'' species group within the genus ''Cebus,'' which also includes the Colombian white-faced capuchin, white-fronted capuchin, the weeper capuchin and the Kaapori capuchin.<ref name=groves2/> This genus is also referred to as "gracile" capuchins.
In 2012 a study by Boubli, ''et al.'' demonstrated that ''C. imitator'' and ''C. capucinus'' split up to 2'' ''million years ago.<ref name=boubli>{{cite journal|title=Cebus Phylogenetic Relationships: A Preliminary Reassessment of the Diversity of the Untufted Capuchin Monkeys|author=Boubli, Jean P.|display-authors=etal|journal=American Journal of Primatology|year=2012|pages=1–13|doi=10.1002/ajp.21998|url=http://socgen.ucla.edu/wp-content/uploads/2013/10/CebusPhylogeneticRelationships.pdf|access-date=2018-12-30|pmid=22311697|volume=74|issue=4|s2cid=12171529|archive-date=2016-03-04|archive-url=https://web.archive.org/web/20160304103046/http://socgen.ucla.edu/wp-content/uploads/2013/10/CebusPhylogeneticRelationships.pdf}}</ref><ref name=alfaro>{{cite journal|title=Capuchin Monkey Research Priorities and Urgent Issues|author=Lynch Alfaro, Jessica|display-authors=etal|url=http://socgen.ucla.edu/wp-content/uploads/2014/04/Capuchin-Monkey-Research-Priorities-and-Urgent-Issues.pdf|access-date=2018-12-30|journal=American Journal of Primatology|year=2014|doi=10.1002/ajp.22269|pmid=24668460|pages=1–16|volume=76|issue=8|s2cid=14778572|archive-date=2017-08-10|archive-url=https://web.archive.org/web/20170810115319/http://socgen.ucla.edu/wp-content/uploads/2014/04/Capuchin-Monkey-Research-Priorities-and-Urgent-Issues.pdf}}</ref> Boubli's study also indicated that the Honduran white-faced capuchins, which had previously been considered a to be a possible separate subspecies, ''C. capucinus limitaneus'', was not genetically distinct from the Panamanian white-faced capuchin.<ref name=boubli/>
The Panamanian white-faced capuchin is among the most well-studied capuchin monkey species.<ref name=alfaro/> Even though many previous studies were performed using the scientific name ''C. capucinus'', as of 2014 there had been no field studies of the Colombian white-faced capuchin, so all these studies were of the Panamanian white-faced capuchin.<ref name=alfaro/>
== Physical description == Like other monkeys in the genus ''Cebus'', the Panamanian white-faced capuchin is named after the order of Capuchin friars – the cowls of these friars closely resemble the monkey's head coloration.<ref>{{cite web|title=Capuchin Franciscans F.A.Q.|publisher=Capuchin Franciscans Vocation Office Province of Saint Joseph|url=http://capuchinfranciscans.org/sub_faq.html|access-date=2008-09-01|archive-url=https://web.archive.org/web/20110725135402/http://capuchinfranciscans.org/sub_faq.html|archive-date=2011-07-25}}</ref><ref name=costa>{{cite book|title=The Natural History of Costa Rican Mammals|author=Wainwright, M.|year=2002|pages=135–139|publisher=Zona Tropical|isbn=978-0-9705678-1-9}}</ref> The Panamanian white-faced capuchin has mostly black fur, with white to yellow like fur on the neck, throat, chest, shoulders, and upper arms.<ref name=field>{{cite book|title=Neotropical Rainforest Mammals A Field Guide|edition=Second|author=Emmons, L.|pages=130–131|year=1997|publisher=University of Chicago Press|isbn=978-0-226-20721-6}}</ref> The face is pink or a white-cream color and may have identifying marks such as dark brows or dark fur patches.<ref name=field/><ref>{{cite book|last=Luedtke|first=Karen|title=Jungle Living: A look at life and social behavior of man and monkey in Central American|year=2012|isbn=978-0-9832448-2-0|pages=40–45}}</ref><ref>{{cite book|title=Jungle Living: A look at life and social behavior of man and monkey in central america|author=Luedtke, K.|page=45|year=2012|isbn=978-0-9832448-2-0}}</ref> An area of black fur on the crown of the head is distinctive.<ref name=field/><ref name=pic>{{cite book|title=The Pictorial Guide to the Living Primates|url=https://archive.org/details/pictorialguideto0000rowe|url-access=registration|author=Rowe, N.|page=[https://archive.org/details/pictorialguideto0000rowe/page/95 95]|year=1996|publisher=Pogonias Press|isbn=978-0-9648825-0-8}}</ref> It has a prehensile tail that is often held coiled, giving the white-faced capuchins the nickname "ringtail".<ref name=field/><ref>{{cite web|title=Medical Dictionary Capuchin Monkey|url=http://medical-dictionary.thefreedictionary.com/Capuchin+(monkey)|publisher=Farlex Inc.|access-date=2008-09-01}}</ref>
Adults reach a length of between {{convert|335|and|453|mm|in|frac=8|abbr=on}}, excluding tail, and a weight of up to {{convert|3.9|kg|lboz|abbr=on}}.<ref name=field/><ref name=pic/> The tail is longer than the body, at up to {{convert|551|mm|in|frac=8|abbr=on}} in length.<ref name=field/><ref name=pic/> Males are about 27% larger than females.<ref name=perspective>{{cite book|title=Primates in Perspective|chapter=The Cebines|author=Jack, K.|editor = Campbell, C. |editor2=Fuentes, A. |editor3=MacKinnon, K. |editor4=Panger, M. |editor5=Bearder, S|year=2007|pages=107–120|publisher=Oxford University Press|isbn=978-0-19-517133-4}}</ref> The brain of a white-faced capuchin is about {{convert|79.2|g|oz|frac=16|abbr=on}}, which is larger than that of several larger monkey species, such as the mantled howler.<ref name=pic/><ref>{{cite book|title=The Pictorial Guide to the Living Primates|url=https://archive.org/details/pictorialguideto0000rowe|url-access=registration|author=Rowe, N.|page=[https://archive.org/details/pictorialguideto0000rowe/page/109 109]|year=1996|publisher=Pogonias Press|isbn=978-0-9648825-0-8}}</ref>
The Panamanian white-faced capuchin is similar to the Colombian white-faced capuchin in appearance, except that the female Panamanian white-faced capuchins have brownish or grayish elongated frontal tufts, which provide a contrast to the pure white cheeks and throat.<ref name=handbook/><ref name=all>{{cite book|title=All the World's Primates|author1=Melin, Amanda D. |author2=Jack, Katherine M. |author3=Fedigan, Linda |author4=Mendez-Carvajal |author5=Pedro G. |editor=Rowe, Noel |editor2=Myers, Marc |pages=286–288|year=2016|publisher=Pogonias Press|isbn=978-1-940496-06-1}}</ref>
== Behavior ==
=== Social structure === thumb|Walking on four limbs The Panamanian white-faced capuchin is a diurnal and arboreal animal.<ref name=field/> However, it does come down to the ground more often than many other New World monkeys.<ref name=ethology>{{cite book|title=Primate Ethology|author1=Morris, D. |author2=Bruce, D. |name-list-style=amp |pages=237–238|year=2005|isbn=978-0-202-30826-5|publisher=Aldine Transaction}}</ref> It moves primarily by walking on all four limbs.<ref>{{cite book|title=New Perspectives in the Study of Mesoamerican Primates|url=https://archive.org/details/newperspectivess00estr|url-access=limited|chapter=Ontogenetic Influences on Positional Behavior in ''Cebus'' and ''Alouatta''|author=Bezanson, L.|editor = Estrada, A. |editor2=Garber, P. |editor3=Pavelka, M. |editor4=Luecke, L|publisher=Springer|location=New York|year=2006|pages=[https://archive.org/details/newperspectivess00estr/page/n337 333]–344|isbn=978-0-387-25854-6}}</ref> It lives in troops, or groups, of up to 40 monkeys (mean 16, range 4–40)<ref name=complete>{{cite book|title=The Complete Capuchin|author=Fragaszy, D.|author2=Visalberghi, E.|author3=Fedigan, L.|name-list-style=amp|chapter=Life History and Demography|pages=74–79|publisher=Cambridge University Press|year=2004}}</ref> and has a male/female adult sex ratio of 0.71 on average (range 0.54–0.88).<ref name=complete/> With rare exceptions, females spend their entire lives with their female kin.<ref name=complete/><ref>{{cite journal|title=Female dispersal in a female-philopatric species, Cebus capucinus. |author1=Jack, K. |author2=Fedigan, L. |name-list-style=amp |journal=Behaviour|volume=146|issue=4 |pages=471–498|year=2009|doi=10.1163/156853909X404420|citeseerx=10.1.1.619.2612 }}</ref><ref name=lomas>{{cite book | last1=Perry | first1=S. |first2=J. | last2=Manson | title=Manipulative Monkeys: The Capuchins of Lomas Barbudal | location=Cambridge, MA |pages=118, 145–154, 169–214, 229–241| publisher=Harvard University Press | year=2008 | isbn=978-0-674-02664-3}}</ref> Males migrate to new social groups multiple times during the course of their lifetimes, migrating for the first time between 20 months and 11 years of age.<ref name=jack1>{{cite journal|title=Male dispersal patterns in white-faced capuchins, Cebus capucinus Part 1: patterns and causes of natal emigration|author1=Jack, K. |author2=Fedigan, L. |name-list-style=amp |journal=Animal Behaviour|volume=67|issue=4 |pages=761–769|year=2004|doi=10.1016/j.anbehav.2003.04.015|s2cid=53701873 }}</ref><ref name=jack2>{{cite journal|title=Male dispersal patterns in white-faced capuchins, Cebus capucinus Part 2: Patterns and causes of secondary dispersal|author1=Jack, K. |author2=Fedigan, L. |name-list-style=amp |journal=Animal Behaviour|volume=67|issue=4 |pages=771–782|year=2004|doi=10.1016/j.anbehav.2003.06.015|s2cid=10044824 }}</ref> The median age of migration in the Santa Rosa population is 4.5 years.<ref name=jack1/> Males sometimes migrate alone, but more often they migrate in the company of other males who are often their kin.<ref name=lomas/><ref name=jack1/><ref name=jack2/> One of the unusual features of the kinship structure of the Panamanian white-faced capuchin, relative to other primate species, is the high degree of relatedness within groups that results from the long tenures of alpha males who sire most of the offspring.<ref name="Muniz, L., Perry, S., Manson, J., Gilkenson, H., Gros-Louis, J., & Vigilant, L. 2006 156–7">{{cite journal|title=Father-daughter inbreeding avoidance in a wild primate population.|author=Muniz, L.|author2=Perry, S.|author3=Manson, J.|author4=Gilkenson, H.|author5=Gros-Louis, J.|author6=Vigilant, L.|name-list-style=amp|journal=Current Biology|volume=16|pages=156–7|year=2006|doi=10.1016/j.cub.2006.02.055|pmid=16527729|issue=5|bibcode=2006CBio...16.R156M |doi-access=free}}</ref><ref name="Perry, S., Manson, J.H., Muniz, L., Gros-Louis, J., & Vigilant, L. 2008 187–199">{{cite journal|title=Kin-biased Social Behaviour in Wild Adult Female White-faced Capuchins (Cebus capucinus) |author=Perry, S. |author2=Manson, J.H. |author3=Muniz, L. |author4=Gros-Louis, J. |author5=Vigilant, L. |name-list-style=amp|journal=Animal Behaviour|volume=76|pages=187–199|year=2008|doi=10.1016/j.anbehav.2008.01.020|s2cid=53154942 }}</ref> Alpha males have been known to keep their positions as long as 17 years in this species<ref name="Muniz, L., Perry, S., Manson, J., Gilkenson, H., Gros-Louis, J., & Vigilant, L. 2006 156–7"/><ref name="Perry, S., Manson, J.H., Muniz, L., Gros-Louis, J., & Vigilant, L. 2008 187–199"/> and this puts them in the unusual position of being available to sire the offspring of their daughters and granddaughters, who produce their first offspring at about 6–7 years of age.<ref name=complete/><ref name="Perry, S., Manson, J.H., Muniz, L., Gros-Louis, J., & Vigilant, L. 2008 187–199"/> Typically, however, alpha males do not breed with their own daughters, even though they do sire virtually all offspring produced by females unrelated to them.<ref name="Muniz, L., Perry, S., Manson, J., Gilkenson, H., Gros-Louis, J., & Vigilant, L. 2006 156–7"/> Those subordinate males who are allies of the alpha male in group defense are the males who sire the offspring of the alpha male's daughters. The high degree to which alpha males monopolize matings results in an unusually large number of paternal half-siblings and full siblings in this species relative to other primate species.<ref name="Perry, S., Manson, J.H., Muniz, L., Gros-Louis, J., & Vigilant, L. 2008 187–199"/>
Kinship is an important organizing factor in the structuring of female-female social relationships.<ref name="Perry, S., Manson, J.H., Muniz, L., Gros-Louis, J., & Vigilant, L. 2008 187–199"/> Particularly in larger groups, females preferentially associate with, groom, and provide coalitionary support to their matrilineally related female kin. They do not exhibit a similar preference for their paternal half sisters, which may mean that they only are capable of recognizing kinship through the maternal line.<ref name="Perry, S., Manson, J.H., Muniz, L., Gros-Louis, J., & Vigilant, L. 2008 187–199"/> Dominance rank is also an important organizing factor, with females more often grooming and associating with females who are closer to them in the dominance hierarchy.<ref name="Perry, S., Manson, J.H., Muniz, L., Gros-Louis, J., & Vigilant, L. 2008 187–199"/> Female-female dyads groom far more than male-female and male-male dyads.<ref name="Perry, S. 1996 167–182">{{cite journal|title=Female-female relationships in wild white-faced capuchin monkeys, Cebus capucinus.|author=Perry, S.|journal=American Journal of Primatology|volume=40|issue=2|pages=167–182|year=1996|doi=10.1002/(SICI)1098-2345(1996)40:2<167::AID-AJP4>3.0.CO;2-W|url=https://deepblue.lib.umich.edu/bitstream/2027.42/38432/1/4_ftp.pdf|hdl=2027.42/38432|s2cid=37033722 |hdl-access=free}}</ref> Coalitionary aggression is common both among males and females, and capuchins seem to have an excellent understanding of the alliance structure in their group. For example, when capuchins are fighting, they sensibly recruit aid from someone who is both higher ranking than they are and also better friends with themselves than with their opponent.<ref>{{cite journal|title=White-faced capuchin monkeys exhibit triadic awareness in their choice of allies.|author=Perry, S.|author2=Manson, J.|author3=Barrett, H.C.|name-list-style=amp|journal=Animal Behaviour|volume=67|pages=165–170|year=2004|doi=10.1016/j.anbehav.2003.04.005|s2cid=53198039}}</ref>
Female capuchins have linear dominance hierarchies.<ref name="Perry, S. 1996 167–182"/><ref>{{cite journal|title=Strength and stability of dominance hierarchies in female white-faced capuchins (Cebus capucinus) at Santa Rosa National Park, Costa Rica. |author1=Bergstrom, M. |author2=Fedigan, L.M. |name-list-style=amp |journal=American Journal of Primatology |volume=71|issue=Suppl 1|page=103|year=2009}}</ref> In contrast to many Old World monkeys such as macaques, in which females socially inherit the rank just below their mothers and just above their next oldest sisters, capuchins do not have a highly predictable ranking within their matrilines.<ref name="Perry, S., Manson, J.H., Muniz, L., Gros-Louis, J., & Vigilant, L. 2008 187–199"/> Males are typically dominant to females.<ref name=Perry-1997>{{cite journal|title=Male-female social relationships in wild white-faced capuchin monkeys, Cebus capucinus.|author=Perry, S.|journal=Behaviour|volume=134|issue=7|pages=477–510|year=1997|doi=10.1163/156853997X00494}}</ref> The alpha male is always easy to discern, but there are sometimes ambiguous rankings among subordinate males.<ref name=lomas/><ref name="Perry, S. 1998 1–34">{{cite journal|title=Male-male social relationships in wild white-faced capuchins, Cebus capucinus. |author=Perry, S.|journal=Behaviour |volume=135|issue=2|pages=1–34|year=1998|doi=10.1163/156853998793066384}}</ref> Male-male relationships are tense, and affiliation between males is typically expressed by resting in contact, playing, or non-conceptive sex rather than by grooming.<ref name="Perry, S. 1998 1–34"/><ref>{{cite journal|title=Nonconceptive sexual behavior in bonobos and capuchins.|author= Manson, J.H.|author2= Perry, S.|author3= Parish, A.R.|name-list-style=amp|journal=International Journal of Primatology| volume=18|issue= 5|pages=767–786|year=1997|doi=10.1023/A:1026395829818|s2cid= 3032455}}</ref> Males cooperate in coalitions against potential predators, and also in defense of the group against other males.<ref name=lomas/><ref name="Perry, S. 1998 1–34"/><ref name="Perry, S. 1996 309–330"/> Occasionally male coalitionary aggression becomes so violent that males are killed, particularly if they are encountered roaming the forest unaccompanied by allies.<ref name=lomas/><ref name="Gros-Louis, J., Perry, S. & Manson, J.H. 2003 341–346">{{cite journal|title=Violent coalitionary attacks and intraspecific killing in wild capuchin monkeys (Cebus capucinus).|author=Gros-Louis, J.|author2=Perry, S.|author3=Manson, J.H.|name-list-style=amp|journal=Primates|volume=44|pages=341–346|year=2003|doi=10.1007/s10329-003-0050-z|pmid=12910384|issue=4|s2cid=6573597}}</ref> Because aggression from other male capuchins is the leading cause of death (aside from poaching by humans, where there is contact between humans and capuchins), male allies are critical for self-defense during migration, and to assist in taking over other groups.<ref name=lomas/> Male emigration to a new troop typically occurs about every 4 years, so most males are in constant danger of having to defend themselves against other groups of males.<ref name=costa/><ref name=male>{{cite journal|title=The Demographic and Reproductive Context of Male Replacements in Cebus Capucinus|author1=Fedigan, L. |author2=Jack, K. |name-list-style=amp |journal=Behaviour|volume=141|issue=6 |pages=755–775|year=2004|url=https://www.ucalgary.ca/~fedigan/Beh%20Fedigan%20and%20Jack%202004.pdf|access-date=2008-11-14|doi=10.1163/1568539042245178}}</ref><ref name=tax2>{{cite book|title=New Perspectives in the Study of Mesoamerican Primates|url=https://archive.org/details/newperspectivess00estr|url-access=limited|chapter=Dominance and Reproductive Success in Wild White-Faced Capuchins|author1=Jack, K. |author2=Fedigan, L. |name-list-style=amp |editor = Estrada, A. |editor2=Garber, P. |editor3=Pavelka, M. |editor4=Luecke, L|publisher=Springer|location=New York|year=2006|pages=[https://archive.org/details/newperspectivess00estr/page/n370 367]–382|isbn=978-0-387-25854-6}}</ref><ref name=":0">{{Cite journal|title = Why Be Alpha Male?|author = Jack|author2 = Fedigan|name-list-style=amp|date = 2006|journal = New Perspectives in the Study of Mesoamerican Primates|doi = 10.1007/0-387-25872-8_18}}</ref>
Immigrating males often kill young infants when they take over a group.<ref name=lomas/><ref>{{cite journal|title=Three apparent cases of infanticide by males in wild white-faced capuchins (Cebus capucinus) |vauthors=Manson JH, Gros-Louis J, Perry S | journal=Folia Primatologica|volume=75|pages=104–106|year=2004|doi=10.1159/000076270|pmid=15010584|issue=2|s2cid=35102249 }}</ref><ref name="Fedigan, L.M. 2003 723–741">{{cite journal| title=Impact of male takeovers on infant deaths, births, and conceptions in Cebus capucinus at Santa Rosa, Costa Rica.|author=Fedigan, L.M.|journal=International Journal of Primatology|volume= 24|issue=4|pages=723–741|year=2003| doi=10.1023/A:1024620620454|s2cid=19279800}}</ref> Females band together to defend their infants from infanticidal males, but they rarely succeed in saving their infants.<ref name=lomas/> Because infants inhibit their mothers from ovulating by nursing frequently, males are able to bring females into estrus earlier by killing the infants and thereby terminating nursing; this has the effect of increasing their breeding opportunities.<ref name=lomas/><ref name="Fedigan, L.M. 2003 723–741"/><ref>{{cite journal|title=Male-male competition and infanticide among the langurs (Presbytis entellus) of Abu, Rajasthan. |author=Hrdy, S.|journal=Folia Primatologica|volume=22|issue=1|pages=19–58|year=1974|doi=10.1159/000155616|pmid=4215710}}</ref> Females do often mate with the killers of their infants, and with time, they typically become as supportive of the new alpha male as they had been of the previous one.<ref name=lomas/> The alpha male helps defend females from subordinate males within the group as well as from infanticidal males from other groups.<ref name=lomas/><ref name=Perry-1997/>
=== Interactions between groups ===
Panamanian white-faced capuchin troops occupy home ranges of between {{convert|32|and|86|ha|acre}}.<ref name=pic/> They travel between {{convert|1|and|3|km|mi|frac=4|abbr=on}} daily, averaging {{convert|2|km|mi|frac=4|abbr=on}} per day.<ref name=colombia>{{cite book|title=Primates of Colombia|author=Defler, T.|pages=227–235|year=2004|isbn=978-1-881173-83-0|publisher=Conservation International|location=Bogotá, D.C., Colombia}}</ref> Although they engage in activity that has been described as "territorial", more recent research indicates that white-faced capuchin troops tend to behave aggressively to other white-faced capuchin troops regardless of where they meet, and the aggression is not necessarily intended to exclude the other troops from a specific home range.<ref>{{cite book|title=The Complete Capuchin|chapter=Behavioral Ecology|author=Fragaszy, D.|author2=Visalberghi, E.|author3=Fedigan, L.|name-list-style=amp|year=2004|pages=38–39|publisher=Cambridge University Press|isbn=978-0-521-66768-5}}</ref>
Home ranges overlap extensively,<ref name="Perry, S. 1996 309–330">{{cite journal|title=Intergroup encounters in wild white-faced capuchins, Cebus capucinus.|author=Perry, S.|journal=International Journal of Primatology|volume=17|issue=3|pages=309–330|year=1996|doi=10.1007/BF02736624|s2cid=20334207}}</ref><ref name="Crofoot, M.C., Gilby, I.C., Wikelski, M.C,& Kays, RW 2008 577–581">{{cite journal|title=Interaction location outweighs the competitive advantage of numerical superiority in Cebus capucinus intergroup contests.|author=Crofoot, M.C.|author-link= Margaret Crofoot |author2=Gilby, I.C.|author3=Wikelski, M.C|author4=Kays, RW|name-list-style=amp|journal=PNAS |volume=105|pages=577–581|year=2008|doi=10.1073/pnas.0707749105|pmid=18184811|issue=2|pmc=2206578|bibcode=2008PNAS..105..577C|doi-access=free}}</ref> so groups are not territorial in the strictest sense of the word. Perhaps because of the intensity of male-male competition and the threat of infanticide, interactions between groups are typically hostile: the males display aggressively toward one another and sometimes engage in physical aggression (even killing an opponent), while females grab their infants and run.<ref name="Perry, S. 1996 309–330"/><ref name="Gros-Louis, J., Perry, S. & Manson, J.H. 2003 341–346"/> Typically, males are the primary participants in aggressive intergroup encounters, and it seems likely that males are defending access to the females in their groups.<ref name="Perry, S. 1996 309–330"/> Alpha males, who have the largest reproductive stake in the group, participate at a higher rate than subordinate males.<ref name="Perry, S. 1996 309–330"/> Groups with more males have an advantage over groups with fewer males, but the location of the encounter within the home range matters as well; smaller groups defeat larger groups when the contest occurs in the core or center area of the smaller group's home range.<ref name="Crofoot, M.C., Gilby, I.C., Wikelski, M.C,& Kays, RW 2008 577–581"/>
=== Interspecific interactions ===
The Panamanian white-faced capuchin sometimes interacts with other sympatric monkey species. Panamanian white-faced capuchins sometimes travel with and even groom Geoffroy's spider monkeys.<ref name=pic/><ref name=colombia/> However, aggressive interactions between the capuchins and spider monkeys also occur.<ref name=interact>{{cite journal|title=Interspecific Interactions between ''Cebus capucinus'' and other Species: Data from Three Costa Rican Sites|author=Rose, L.|author2=Perry, S.|author3=Panger, M.|author4=Jack, K.|author5=Manson, J.|author6=Gros-Louis, J.|author7=Mackinnin, K.|name-list-style=amp|journal=International Journal of Primatology|volume=24|issue=4|date=August 2003|url=http://people.ucsc.edu/~evogel/cv/Roseetal2003.pdf|pages=780–785|doi=10.1023/A:1024624721363|s2cid=430769|access-date=2008-09-04|archive-url=https://web.archive.org/web/20090225160139/http://people.ucsc.edu/~evogel/cv/Roseetal2003.pdf|archive-date=2009-02-25}}</ref> Interactions between the Panamanian white-faced capuchin and mantled howler are infrequent, and sometimes result in the capuchins threatening the larger howlers.<ref name=colombia/> However, affiliative associations between the capuchins and howlers do sometimes occur, mostly involving juveniles playing together.<ref name=interact/>
Although South American capuchin species often travel with and feed together with squirrel monkeys, the Panamanian white-faced capuchin only rarely associates with the Central American squirrel monkey. This appears to be related to the patchier, more dispersed distribution of food resources in Central America and the fact that there is less dietary overlap between the Central American squirrel monkey and the white-faced capuchin than between their South American counterparts. Therefore, there is less benefit to the Central American squirrel monkey in associating with the Panamanian white-faced capuchin in order to exploit the capuchin's knowledge of food resource distribution. In addition, compared to their South American counterparts, male Panamanian white-headed capuchins are relatively more alert to rival males than to predators, reducing the predator detection benefits that the Central American squirrel monkey receives from associating with the Panamanian white-faced capuchin compared to its South American counterparts. Since the squirrel monkeys generally initiate interactions with the capuchins in South America, the fact that similar associations would impose higher foraging costs and impart fewer predator detection benefits to the Central American squirrel monkey leads to fewer associations with the Panamanian white-faced capuchin.<ref name=perspective/><ref name=move>{{cite book|title=On the Move|chapter=Social Manipulation Within and Between Troops Mediates Primate Group Movement|author=Boinski, S.|editor=Boinski, S. |editor2=Garber, P.|year=2000|publisher=The University of Chicago Press|isbn=978-0-226-06340-9|pages=447–448}}</ref><ref>{{cite journal|title=Why don't ''Saimiri oerstedii'' and ''Cebus capucinus'' form mixed-species groups?|author=Boinski, S.|journal=International Journal of Primatology|volume=10|issue=2|date=April 1989|pages=103–114|doi=10.1007/BF02736248|s2cid=24192169|url=https://zenodo.org/record/1232560|type=Submitted manuscript}}</ref><ref>{{cite book|title=The Complete Capuchin|chapter=Community Ecology|author=Fragaszy, D.|author2=Visalberghi, E.|author3=Fedigan, L.|name-list-style=amp|year=2004|page=70|publisher=Cambridge University Press|isbn=978-0-521-66768-5}}</ref>
On the island of Jicarón, Panamanian white-faced capuchins coexist with Coiba Island howler monkeys (''Alouatta coibensis coibensis'').<ref>{{MSW3 Groves|pages=148–149|id=12100377}}</ref> In early 2022, motion-triggered cameras detected male capuchins carrying infant howlers on their backs while walking or smashing seeds with rocks. Further research indicated that during 2022 and 2023, at least five male capuchins abducted at least 11 infant howler monkeys and carried them on their backs. The capuchins were unable to properly care for or feed the infants, and at least four—and probably most or all—of the baby howlers died. By the time they announced their findings in May 2025, researchers had concluded that the cameras had captured a rare behavior that had arisen as a "fashion fad" among a single population of male capuchins, and described the capuchins' behavior as "the first known documentation of a social tradition in which animals repeatedly abduct and carry infants of another species—without any clear benefit to themselves."<ref>{{cite web|author=<!--Not stated--> |url=https://www.mpg.de/24686977/0507-ornr-capuchin-monkeys-develop-bizarre-fad-of-abducting-baby-howlers-987453-x |title=Capuchin monkeys caught abducting baby howlers: Cameras on a remote island in Panama captured the origin and spread of a bizarre "fad" |website=Max-Planck-Gesellschaft|publisher=Max Planck Institute|date=19 May 2025|access-date=1 June 2025}}</ref><ref>{{cite journal |last1=Goldsborough |first1=Zoë |last2=Crofoot |first2=Margaret C. |last3=Jacobson |first3=Odd T. |last4=Corewyn |first4=Lisa |last5=Rosario-Vargas |first5=Eveyn del |last6=León |first6=Julián |last7=Barrett |first7=Brendan J. |date=19 May 2025 |title=Rise and spread of a social tradition of interspecies abduction |url=https://www.cell.com/current-biology/fulltext/S0960-9822(25)00372-0 |journal=Current Biology |volume= 35|issue= 10|pages=375–376 |doi=10.1016/j.cub.2025.03.056 |pmid=40393397 |access-date=1 June 2025|doi-access=free }}</ref><ref>{{cite news|last=Larson |first=Christina |url=https://www.washingtonpost.com/politics/2025/05/19/monkey-kidnappings-panama/cfcca8cc-34c2-11f0-9c9e-0db2d748bea7_story.html |title=Caught on camera, capuchin monkeys kidnap howler monkey babies: Scientists have found evidence of monkey kidnappings in Panama |newspaper=The Washington Post|date=19 May 2025|access-date=1 June 2025}}</ref>
Several non-primate animal species tend to follow troops of white-faced monkeys or are otherwise attracted by their presence. White-lipped peccaries and agoutis are attracted by feeding white-faced capuchins, looking for fruit that the capuchins drop.<ref name=colombia/> Several species of bird are also known to follow Panmanian white-faced capuchins looking for food. These include the double-toothed kite, the white hawk and the sharp-shinned hawk.<ref name=colombia/>
=== Diet === [[File:White faced Capuchin.jpg|right|thumb|White-faced capuchin eating a wild banana along the Frío River, Costa Rica]] thumb|left|Foraging in the trees The Panamanian white-faced capuchin is an omnivore. Its primary foods are fruit and insects.<ref name=costa/> It forages at all levels of the forest, including the ground.<ref name=colombia/> Methods for finding food include stripping bark off of trees, searching through leaf litter, breaking dead tree branches, rolling over rocks, and using stones as anvils to crack hard fruits.<ref name=chapman/> Its prehensile tail assists with feeding, helping support the monkey when foraging for food below the branches.<ref name=colombia/>
Fruit can make up between 50% and 67% or more of the capuchin's diet.<ref name=costa/> In one study in Panama, white-faced capuchins ate 95 different fruit species.<ref name=costa/> Among its favorite fruits are figs from the family Moraceae, mangos and related fruits from the family Anacardiaceae, the bean-like fruits from the family Leguminosae and fruits from the family Rubiaceae.<ref name=diet>{{cite book|title=The Complete Capuchin|chapter=Behavioral Ecology|author=Fragaszy, D.|author2=Visalberghi, E.|author3=Fedigan, L.|name-list-style=amp|year=2004|pages=43–47|publisher=Cambridge University Press|isbn=978-0-521-66768-5}}</ref> It will also eat fruits from Euphorbiaceae such as Mexican jumping bean ''Sebastiania pavoniana''.<ref name="Melin">{{cite journal |last1=Melin |first1=Amanda D. |last2=Fedigan |first2=Linda Marie |last3=Hiramatsu |first3=Chihiro |last4=Kawamura |first4=Shoji |title=Polymorphic color vision in white-faced capuchins (Cebus capucinus): Is there foraging niche divergence among phenotypes? |journal=Behavioral Ecology and Sociobiology |date=22 September 2007 |volume=62 |issue=5 |page=663 |doi=10.1007/s00265-007-0490-3 |s2cid=13839857 |url=https://www.researchgate.net/publication/225902185 |access-date=20 July 2018 |language=en |format=PDF |issn=1432-0762 |oclc=437741616}}</ref> It generally only eats ripe fruit, testing for ripeness by smelling, tasting and prodding the fruit.<ref name=costa/> It typically eats only the pulp and juice, spitting out the seeds and fibers.<ref name=costa/> Other plant matter eaten includes flowers, young leaves, seeds of certain plants, and bromeliads.<ref name=costa/><ref>{{cite book|title=New Perspectives in the Study of Mesoamerican Primates|url=https://archive.org/details/newperspectivess00estr|url-access=limited|chapter=Food Choice by Juevenile Capuchin Monkeys|author=MacKinnon, K.|editor = Estrada, A. |editor2=Garber, P. |editor3=Pavelka, M. |editor4=Luecke, L|publisher=Springer|location=New York|year=2006|pages=[https://archive.org/details/newperspectivess00estr/page/n358 354]–360|isbn=978-0-387-25854-6}}</ref> It also uses the bromeliads as a water source, drinking the water that gets trapped inside.<ref name=costa/> In Carara National Park the capuchins have a varied diet in addition to the above of banana fruits and flowers, heliconia seeds, huevos de caballo fruits and anacardiaceae stems.<ref>{{cite book|last=Luedtke|first=Karen|title=Costa Rica: Monkeys, Animal Behavior, Cognitive Neuroscience|year=2010|chapter=Ch 11|isbn=978-0-9832448-0-6|page=109}}</ref>
Insect prey eaten includes beetle larvae, butterfly and moth caterpillars, ants, wasps, and ant and wasp larvae.<ref name=costa/> It also eats larger prey, such as birds, bird eggs, frogs, lizards, crabs, mollusks and small mammals.<ref name=costa/><ref name=life>{{cite video|title=Life of Mammals|people=David Attenborough|date=2003|publisher=BBC Video}}</ref> The population in Guanacaste, Costa Rica, in particular is noted for hunting squirrels, magpies, white-crowned parrots<ref name=costa/> and baby coatis.<ref>{{cite journal | author = Perry S. Rose L. | year = 1994 | title = Begging and transfer of coati meat by white-faced capuchin monkeys, Cebus capucinus | url = http://deepblue.lib.umich.edu/bitstream/2027.42/41610/1/10329_2006_Article_BF02381950.pdf | journal = Primates | volume = 35 | issue = 4| pages = 409–415 | doi=10.1007/bf02381950| hdl = 2027.42/41610 | s2cid = 146496 | hdl-access = free }}</ref> The amount of vertebrate prey eaten varies by troop.<ref name=costa/> Even neighboring troops can show significant differences in their diets.<ref name=chapman>{{cite journal|title=Dietary Differences between Neighboring ''Cebus capucinus'' Groups: Local Traditions, Food Availability or Responses to Food Profitability?|author1=Chapman, C. |author2=Fedigan, L. |name-list-style=amp |journal=Folia Primatol|volume=54|pages=177–186|year=1990|url=http://www.arts.mcgill.ca/programs/anthro/chapman_files/CWeb/Pdf/33_CRNeighboringCebus.pdf|doi=10.1159/000156442|pmid=2391047|issue=3–4}}</ref>
The diet can vary between the rainy and dry season. For example, in Guanacaste, Costa Rica, the Panamanian white-faced capuchin can eat a wide variety of fruits as well as caterpillars in the early rainy season (June to November).<ref name=diet/> But during the dry season, only figs and a few other types of fruit are available.<ref name=diet/> During the dry season, chitinous insects, ant and wasp larvae and vertebrates become a particularly important part of the Panamanian white-faced capuchin's diet.<ref name=diet/> Access to water can also become an issue during the dry season. The Panamanian white-faced capuchin likes to drink daily, so in forests where water holes dry up during the dry season, there can be competition between troops over access to the remaining water holes.<ref name=diet/>
=== Tool use ===
Capuchins are considered among the most intelligent of the New World monkeys; they have been the subject of many studies on behaviour and intelligence. The capuchins' intelligence is thought to be an adaptation to support their feeding habits; they rely on ephemeral food sources which may be hard to find. In one particular study conducted in 2007, capuchins were found to be among the ten most intelligent primates, second to spider monkeys among New World monkeys.<ref name=intelligence>{{cite news|title=Chimps Knocked Off Top of the IQ Tree|url=http://www.timesonline.co.uk/tol/news/uk/article1654998.ece|archive-url=https://web.archive.org/web/20070515174102/http://www.timesonline.co.uk/tol/news/uk/article1654998.ece|archive-date=May 15, 2007|author1=Leake, D. |author2=Dobson, R. |name-list-style=amp |date=April 15, 2007|access-date=2008-09-01 | location=London|work=The Times}}</ref>
The use of stone tools is a marked difference between the gracile capuchins of the genus ''Cebus'' and the robust capuchins of the genus ''Sapajus''. Although widespread in robust capuchins, only one case of habitual stone tool use has been reported by gracile capuchins. One population of Panamanian white-faced capuchins found in Coiba National Park in Panama has been observed using hammerstones and anvils to process fruits from ''Terminalia catappa, Bactris major'', and ''Cocos nucifera'' (coconuts) and invertebrates such as nerite snails, hermit crabs, and Halloween crabs.<ref name="Barrett_etal2018">{{Cite journal|last1=Barrett|first1=Brendan J.|last2=Monteza-Moreno|first2=Claudio M.|last3=Dogandžić|first3=Tamara|last4=Zwyns|first4=Nicolas|last5=Ibáñez|first5=Alicia|last6=Crofoot|first6=Margaret C.|title=Habitual stone-tool-aided extractive foraging in white-faced capuchins, ''Cebus capucinus''|doi-access=free|journal=Royal Society Open Science|year=2018|volume=5|issue=8|article-number=181002|doi=10.1098/rsos.181002|pmc=6124021|pmid=30225086|bibcode=2018RSOS....581002B}}</ref><ref>{{Cite journal|last1=Monteza-Moreno|first1=Claudio M.|last2=Dogandžić|first2=Tamara|last3=McLean|first3=Kevin A.|last4=Castillo-Caballero|first4=Pedro L.|last5=Mijango-Ramos|first5=Zarluis|last6=Del Rosario-Vargas|first6=Evelyn|last7=Crofoot|first7=Margaret C.|last8=Barrett|first8=Brendan J.|date=2020-06-01|title=White-Faced Capuchin, ''Cebus capucinus imitator'', Hammerstone and Anvil Tool Use in Riparian Habitats on Coiba Island, Panama|journal=International Journal of Primatology|language=en|volume=41|issue=3|pages=429–433|doi=10.1007/s10764-020-00156-5|s2cid=218773276|issn=1573-8604|url=https://kops.uni-konstanz.de/bitstreams/2d86f190-0c91-4724-ae17-7d4c4d17912b/download}}</ref> The Panamanian white-faced capuchin is known to rub parts of certain plants into their hair. Plants used in this manner include citrus fruits, vines of the genera ''Piper'' and ''Clematis'', monkey comb (genus ''Sloanea''), dumb cane and custard apple.<ref name=costa/><ref name=life/> Ants and millipedes are also used in this way.<ref name=costa/> It is not definitively known what this rubbing is for, but this may deter parasites such as ticks and insects, or it may serve as a fungicide or bactericide or anti-inflammatory agent.<ref name=costa/> Alternatively, it may be a form of scent marking.<ref name=costa/> The Panamanian white-faced capuchin also uses tools in other ways. It has been known to beat snakes with sticks in order to protect itself or to get the snake to release an infant,<ref name=costa/> and it sometimes uses sticks as probes to explore openings.<ref>{{cite book|title=The Complete Capuchin|chapter=Capuchins Use Objects as Tools|author=Fragaszy, D.|author2=Visalberghi, E.|author3=Fedigan, L.|name-list-style=amp|year=2004|pages=173–183|publisher=Cambridge University Press|isbn=978-0-521-66768-5}}</ref> In captivity, it has been known to use tools to get to food or to defend itself, and in one case a white-faced capuchin used a squirrel monkey as a projectile, hurling it at a human observer.<ref name=costa/>
The Panamanian white-faced capuchin's intelligence and ability to use tools allows it to be trained to assist paraplegics.<ref name=quad>{{cite web|title=Capuchin Monkey (''Cebus capucinus'')|url=http://www.rainforest-alliance.org/resources.cfm?id=capuchin_monkey|publisher=The Rainforest Alliance|access-date=2009-02-07}}</ref> Other species of capuchin monkeys are also trained in this manner.<ref>{{cite news|title=Monkeys as Helpers To Quadriplegics At Home |last=Blumenthal|first=D.|date=June 17, 1987|work=The New York Times}}</ref> Panamanian white-faced capuchins can also be trained for roles on television and movies, such as Marcel on the television series ''Friends''.<ref>{{cite news|title=Earth Matters: Turkey struggles with national epidemic: primate smuggling|last=Pflum|first=M.|publisher=CNN|date=March 18, 2000|url=http://www.monkeyworld.co.uk/press.php?ArticleID=15|access-date=2009-02-07|archive-date=2006-09-28|archive-url=https://web.archive.org/web/20060928022225/http://www.monkeyworld.co.uk/press.php?ArticleID=15}}</ref> They were also traditionally used as organ grinder monkeys.<ref name=henderson/>
=== Communication === thumb|right|Facial expression
The Panamanian white-faced capuchin is noisy.<ref name=field/> Loud calls, such as barks and coughs, are used to communicate threat warnings, and softer calls, such as squeals, are used in intimate discourse.<ref name=costa/> Different types of threats, such as a threat from a terrestrial animal versus a threat from a bird, invoke different vocalizations.<ref name=colombia/> Facial expressions and scent are also important to communication.<ref name=groom>{{cite book|title=The Complete Capuchin|chapter=Social Interactions, Relationships and Social Structure|author=Fragaszy, D.|author2=Visalberghi, E.|author3=Fedigan, L.|name-list-style=amp|year=2004|pages=202–220|publisher=Cambridge University Press|isbn=978-0-521-66768-5}}</ref> It sometimes engages in a practice known as "urine washing", in which the monkey rubs urine on its feet.<ref>{{cite book|title=The Complete Capuchin|chapter=The Body|author=Fragaszy, D.|author2=Visalberghi, E.|author3=Fedigan, L.|name-list-style=amp|year=2004|page=102|publisher=Cambridge University Press|isbn=978-0-521-66768-5}}</ref> The exact purpose of this practice is unknown, but it may be a form of olfactory signal.<ref name=groom/>
== Reproduction == [[File:Young White-headed capuchin.JPG|thumb|Juvenile in Palo Verde National Park, Costa Rica]] The Panamanian white-faced capuchin uses a polygamous mating system in which a male may mate with multiple females.<ref name=colombia/> Although the dominant male does not monopolize breeding, studies have shown that the dominant male does tend to father most of the young.<ref name=tax2/> Although a female may mate with several males, the dominant male may be more likely to copulate when the female is at peak fertility.<ref name=tax2/><ref>{{cite book|title=New Perspectives in the Study of Mesoamerican Primates|url=https://archive.org/details/newperspectivess00estr|url-access=limited|chapter=Post-conceptive Mating in White-Faced Capuchins|author=Carnegie, S.|author2=Fedigan, L.|author3=Ziegler, T.|name-list-style=amp|editor = Estrada, A. |editor2=Garber, P. |editor3=Pavelka, M. |editor4=Luecke, L|publisher=Springer|location=New York|year=2006|pages=[https://archive.org/details/newperspectivess00estr/page/n390 387]–405|isbn=978-0-387-25854-6}}</ref> Nonetheless, there is evidence that dominant males do tend to avoid breeding with their own daughters who are members of the troop.<ref name=sa>{{cite book|title=South American Primates: Comparative Perspectives in the Study of Behavior, Ecology and Conservation|chapter=Genetic Approaches to the Study of Dispersal and Kinship in New World Primates|author=Di Fiore, A.|editor=Garber, P. |editor2=Estrada, A. |editor3=Bicca-Marques, J.C. |editor4=Heymann, E. |editor5=Strier, K|editor5-link=Karen B. Strier|pages=222–223|year=2009|publisher=Springer|isbn=978-0-387-78704-6}}</ref> Such avoidance is rare among New World primates.<ref name=sa/>
Copulation takes about 2 minutes, and the gestation period is 5 to 6 months.<ref name=colombia/> Usually a single young is born, but twins occur occasionally. Most births occur during the dry season from December to April.<ref name=pic/><ref name=colombia/> The infant is carried across its mother's back for about 6 weeks.<ref name=colombia/> After about 4 to 5 weeks it can stray from its mother for brief periods and by about 3 months it can move around independently, although some infants will be mostly independent earlier. Weaning occurs between 6 and 12 months. While the mother rests, the young spends most of its time foraging or playing, either on its own or with other juveniles.<ref name=colombia/> Capuchins engage in high levels of alloparenting, in which monkeys other than the mother help care for the infant.<ref name=forage>{{cite journal|author=Panger, M.|author2=Perry, S.|author3=Rose, L.|author4=Gros-Louis, J.|author5=Vogel, E.|author6=Mackinnon, C.|author7=Baker, M.|name-list-style=amp|title=Cross-Site Differences in Foraging Behavior of White-Faced Capuchins (''Cebus capucinus'')|url=http://people.ucsc.edu/~njdominy/people/pdf/Panger_et_al.pdf|journal=American Journal of Physical Anthropology|volume=119|issue=1|pages=52–66|year=2002|doi=10.1002/ajpa.10103|pmid=12209573|access-date=2009-10-06|archive-date=2011-06-04|archive-url=https://web.archive.org/web/20110604103901/http://people.ucsc.edu/~njdominy/people/pdf/Panger_et_al.pdf}}</ref> Infants are carried by alloparents most often between 4 and 6 weeks in age.<ref name=lomas/> Males as well as females engage in alloparenting.<ref name=lomas/><ref name=colombia/>
Like other capuchin species, the Panamanian white-faced capuchin matures slowly. Sexual maturity can be reached at 3 years.<ref name=henderson/> But on average, females give birth for the first time at 7 years old and give birth every 26 months thereafter.<ref name=perspective/> Males attain reproductive maturity at 10 years old.<ref name=perspective/> The Panamanian white-faced capuchin has a long life span given its size. The maximum recorded life span in captivity is over 54 years.<ref name=perspective/>
== Distribution and habitat == The Panamanian white-faced capuchin is found in much of Central America. In Central America, its range includes much of Honduras, Nicaragua, Costa Rica and Panama.<ref name=handbook/><ref name=tax>{{cite book|title=New Perspectives in the Study of Mesoamerican Primates|url=https://archive.org/details/newperspectivess00estr|url-access=limited|chapter=Taxonomy and Distributions of Mesoamerican Primates|author=Rylands, A.|author2=Groves, C.|author3=Mittermeier, R.|author4=Cortes-Ortiz, L.|author5=Hines, J.|name-list-style=amp|editor = Estrada, A. |editor2=Garber, P. |editor3=Pavelka, M. |editor4=Luecke, L|publisher=Springer|location=New York|year=2006|pages=[https://archive.org/details/newperspectivess00estr/page/n52 40]–43|isbn=978-0-387-25854-6}}</ref> It has also been reported to occur in eastern Guatemala and southern Belize, but these reports are unconfirmed.<ref name=tax/> It is among the most commonly seen monkeys in Central America's national parks, such as Manuel Antonio National Park, Corcovado National Park, Santa Rosa National Park and Soberania National Park.<ref>{{cite book|title=Watching Wildlife Central America|author1=Hunter, L. |author2=Andrew, D. |name-list-style=amp |year=2002|pages=97, 100, 110, 130|publisher=Lonely Planet Publications|isbn=978-1-86450-034-9}}</ref> It appears on the reverse side of the Costa Rican 5,000 colón note.
While the white-faced capuchin is very common in Costa Rica and Panama, the monkey has been largely extirpated from Honduras and much of Nicaragua. Many Honduran capuchins were captured and relocated to the island of Roatán, and many Nicaraguan capuchins were captured and relocated to the island of Ometepe. In Nicaragua, wild capuchins may still be easily spotted in regions around Masaya, as well as around Bluefields and other locations around the South Caribbean coast. They are seen, in the wild, daily by visitors who climb one of the volcanoes on Ometepe Island.<ref>{{Cite web|url=https://ometepeislandinfo.com/Volcan-Maderas|title=Ometepe Island Info - Volcán Maderas|website=ometepeislandinfo.com|language=en|access-date=2017-03-05}}</ref>
It is found in many different types of forest, including mature and secondary forests, and including evergreen and deciduous forests, dry and moist forests, and mangrove and montane forests.<ref name=field/><ref name=henderson>{{cite book|title=Field Guide to the Wildlife of Costa Rica|url=https://archive.org/details/fieldguidetowild00hend|url-access=limited|author=Henderson, C.|year=2000|pages=[https://archive.org/details/fieldguidetowild00hend/page/n400 454]–455|publisher=University of Texas Press|isbn=978-0-292-73459-3}}</ref> However, it appears to prefer primary or advanced secondary forests.<ref name=colombia/> Also, higher densities of white-faced capuchins are found in older areas of forest and in areas containing evergreen forest, as well as areas with more water availability during the dry season.<ref>{{cite book |author1=DeGama, H. |author2=Fedigan, L. |name-list-style=amp | editor = Estrada, A. |editor2=Garber, P. |editor3=Pavelka, M. |editor4=Luecke, L | chapter = The Effects of Forest Fragment Age, Isolation, Size, Habitat Type, and Water Availability on Monkey Density in a Tropical Dry Forest | title = New Perspectives in the Study of Mesoamerican Primates|url=https://archive.org/details/newperspectivess00estr |url-access=limited | pages = [https://archive.org/details/newperspectivess00estr/page/n174 165]–186| publisher=Springer|location=New York| isbn=978-0-387-25854-6 | year = 2006}}</ref>
== Conservation status == [[File:White-faced capuchin monkey 4.jpeg|thumb|left|In Manuel Antonio National Park, Costa Rica]] The Panamanian white-faced capuchin is regarded as vulnerable from a conservation standpoint by IUCN.<ref name=iucn/> It is threatened by deforestation, hunting for pet trade and sometimes for bushmeat and by the fact that farmers sometimes attack them as potential threats.<ref name=iucn/> However, deforestation may also impact its main predator, the harpy eagle, more than it directly impacts the Panamanian white-faced capuchin, and so on a net basis deforestation may not be as harmful to the capuchin's status.<ref name=costa/> The Panamanian white-faced capuchin can adapt to forest fragmentation better than other species due to its ability to live in a wide variety of forest types and exploit a wide variety of food sources.<ref>{{cite book|title=New Perspectives in the Study of Mesoamerican Primates|url=https://archive.org/details/newperspectivess00estr|url-access=limited|chapter=Concluding Comments and Conservation Priorities|author=Garber, P.|author2=Estrada, A.|author3=Pavelka, M.|name-list-style=amp|editor = Estrada, A. |editor2=Garber, P. |editor3=Pavelka, M. |editor4=Luecke, L|publisher=Springer|location=New York|year=2006|pages=[https://archive.org/details/newperspectivess00estr/page/n568 570]–571|isbn=978-0-387-25854-6}}</ref> The Panamanian white-faced capuchin is important to its ecosystems as a seed and pollen disperser.<ref name=costa/><ref name=henderson/> It also impacts the ecosystem by eating insects that act as pests to certain trees, by pruning certain trees, such as ''Gustavia superba'' and ''Bursera simaruba'', causing them to generate more branches and possibly additional fruit, and by accelerating germination of certain seeds when they pass through the capuchin's digestive tract.<ref name=costa/> In addition, the Panamanian white-faced capuchin sometimes kills ''Vachellia collinsii'' plants when it rips through the plant's branches to get to resident ant colonies.<ref name=costa/> {{clear}}
== References == {{Reflist}}
== Further reading == {{refbegin}} *{{cite book |last1=Perry |first1=S. |last2=Manson |first2=J. |year=2009 |chapter=11. Guapo: Innovation and Tradition in the Creation of Bond-Testing Rituals |title=Manipulative Monkeys: The Capuchins of Lomas Barbudal |pages=245–263 |location=Cambridge, MA and London, England |publisher=Harvard University Press}} *{{cite journal |last1=Perry |first1=S. |year=2011 |title=Social traditions and social learning in capuchin monkeys (''Cebus'') |journal=Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences |volume=366 |issue=1567 |pages=988–996 |doi=10.1098/rstb.2010.0317|pmid=21357221 |pmc=3049088 }} *{{cite journal |last1=Perry |first1=S. |last2=Baker |first2=M. |last3=Fedigan |first3=L. |last4=Gros Louis |first4=J. |last5=Jack |first5=K. |last6=MacKinnon |first6=Katherine C. |last7=Manson |first7=Joseph H. |last8=Panger |first8=M. |last9=Pyle |first9=K. |last10=Rose |first10=L. |year=2003 |title=Social Conventions in Wild White-faced Capuchin Monkeys: Evidence for Traditions in a Neotropical Primate |journal=Current Anthropology |volume=44 |issue=2 |pages=241–268 |doi=10.1086/345825|s2cid=54755518 }} *{{cite journal |last1=Perry |first1=S. E. |last2=Barrett |first2=B. J. |last3=Godoy |first3=I. |year=2017 |title=Older, sociable capuchins (''Cebus capucinus'') invent more social behaviors, but younger monkeys innovate more in other contexts |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=114 |issue=30 |pages=7806–7813 |doi=10.1073/pnas.1620739114|pmid=28739946 |pmc=5544268 |bibcode=2017PNAS..114.7806P |doi-access=free }} *{{cite book |last1=Perry |first1=Susan |year=1995 |title=Social relationships in wild white-faced capuchin monkeys, ''Cebus capucinus'' |location=Ann Arbor |publisher=University Microfilms International}} {{refend}}
== External links == *[https://archive.today/20130105063441/http://www3.interscience.wiley.com/journal/110517704/abstract Use of a club by a wild white-faced capuchin to attack a venomous snake] * [http://lagunadeapoyo.blogspot.com/2013/09/animal-rescue-xiv-white-faced-capuchin.html Pet rescue -- white-faced capuchin returned to the wild]
{{Cebidae nav}} {{Costa Rican monkey species}} {{Panamanian monkey species}} {{Taxonbar|from=Q31836365}} {{good article}}
Category:Capuchin monkeys Category:Primates of Central America Category:Least concern biota of North America Category:Tool-using mammals Category:Mammals described in 1903 Category:Taxa named by Oldfield Thomas Category:Species that are or were threatened by deforestation Category:Species that are or were threatened by the pet trade