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Gracile capuchin monkeys are
capuchin monkey 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 fores ...
s 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 monkey 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 ...
s, such as the
tufted capuchin 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 th ...
, and the gracile capuchins. The gracile capuchins retain the genus name ''Cebus'', while the robust species have been transferred to ''
Sapajus 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 ...
''.


Taxonomy

Following Groves (2005), taxa within the genus ''Cebus'' include: *
White-fronted capuchin 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: Bo ...
, ''Cebus albifrons'' **
Ecuadorian capuchin The Ecuadorian capuchin (''Cebus aequatorialis''), or Ecuadorian white-fronted capuchin is a species of gracile capuchin monkey of the family Cebidae. It was formerly classified as a subspecies of the white-fronted capuchin (''C. albifrons'').& ...
, ''Cebus albifrons aequatorialis'' **
Humboldt's white-fronted capuchin Humboldt's white-fronted capuchin (''Cebus albifrons'') is a species of gracile capuchin monkey. It is found in Colombia, Venezuela, Brazil, and potentially the island of Trinidad. Taxonomy The species name ''Cebus albifrons'' was formerly co ...
, ''Cebus albifrons albifrons'' **
Shock-headed capuchin The shock-headed capuchin (''Cebus cuscinus'') is a species of gracile capuchin monkey from Bolivia and Peru. It was previously classified as a subspecies of the Humboldt's white-fronted capuchin (''C. albifrons''), but in 2013 Mittermeier an ...
, ''Cebus albifrons cuscinus'' **
Trinidad white-fronted capuchin The Trinidad white-fronted capuchin is a subspecies (''Cebus albifrons trinitatis'') or species (''Cebus trinitatis'') of gracile capuchin monkey. It is found on the island of Trinidad. Taxonomy Boubli ''et al''. found in a 2012 study that th ...
, ''Cebus albifrons trinitatis'' **
Spix's white-fronted capuchin Spix's white-fronted capuchin (''Cebus unicolor'') is a species of gracile capuchin monkey. It had previously been classified as a subspecies of the Humboldt's white-fronted capuchin (''C. albifrons''). Following genetic studies by Boubli, '' ...
, ''Cebus albifrons unicolor'' **
Varied capuchin The varied white-fronted capuchin (''Cebus versicolor'') is a species of gracile capuchin monkey from Colombia. It had been classified as a subspecies of the white-fronted capuchin (''C. albifrons'') Genetic analysis by Jean Boubli in 2012 reve ...
, ''Cebus albifrons versicolor'' *
White-headed capuchin White-faced capuchin, or white headed capuchin, can refer to either of two species of gracile capuchin monkey: * ''Cebus imitator'', the Panamanian white-faced capuchin, also known as the Panamanian white-headed capuchin or Central American w ...
or white-faced capuchin, ''Cebus capucinus'' *
Kaapori capuchin The Kaapori capuchin (''Cebus kaapori''), 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 populat ...
, ''Cebus kaapori'' *
Wedge-capped capuchin The wedge-capped capuchin or Guianan weeper capuchin (''Cebus olivaceus'') 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 ove ...
, ''Cebus olivaceus'' Subsequent revisions have split some of these into additional species: *
Colombian white-faced capuchin The Colombian white-faced capuchin (''Cebus capucinus''), also known as the Colombian white-headed capuchin or Colombian white-throated capuchin, is a medium-sized New World monkey of the family Cebidae, subfamily Cebinae. It is native to the ext ...
or Colombian white-headed capuchin, ''Cebus capucinus'' *
Panamanian white-faced capuchin 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 for ...
or Panamanian white-headed capuchin, ''Cebus imitator'' *
Marañón white-fronted capuchin The Marañón white-fronted capuchin (''Cebus yuracus'') also or known as Peruvian white-fronted capuchin or Andean white-fronted capuchin is a species of gracile capuchin monkey from the upper Amazon Basin. It had been regarded as synonymous ...
, ''Cebus yuracus'' *
Shock-headed capuchin The shock-headed capuchin (''Cebus cuscinus'') is a species of gracile capuchin monkey from Bolivia and Peru. It was previously classified as a subspecies of the Humboldt's white-fronted capuchin (''C. albifrons''), but in 2013 Mittermeier an ...
, ''Cebus cuscinus'' *
Spix's white-fronted capuchin Spix's white-fronted capuchin (''Cebus unicolor'') is a species of gracile capuchin monkey. It had previously been classified as a subspecies of the Humboldt's white-fronted capuchin (''C. albifrons''). Following genetic studies by Boubli, '' ...
, ''Cebus unicolor'' *
Humboldt's white-fronted capuchin Humboldt's white-fronted capuchin (''Cebus albifrons'') is a species of gracile capuchin monkey. It is found in Colombia, Venezuela, Brazil, and potentially the island of Trinidad. Taxonomy The species name ''Cebus albifrons'' was formerly co ...
, ''Cebus albifrons'' *
Guianan weeper capuchin The wedge-capped capuchin or Guianan weeper capuchin (''Cebus olivaceus'') 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 ...
, ''Cebus olivaceus'' *
Chestnut capuchin The chestnut capuchin or chestnut weeper capuchin (''Cebus castaneus'') is a species of capuchin monkey from northeastern Brazil, southern Guyana, Suriname, and French Guiana. Taxonomy It was described in 1851 as a subspecies of the wedge-ca ...
, ''Cebus castaneus'' *
Ka'apor capuchin The Kaapori capuchin (''Cebus kaapori''), 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 populat ...
, ''Cebus kaapori'' *
Trinidad white-fronted capuchin The Trinidad white-fronted capuchin is a subspecies (''Cebus albifrons trinitatis'') or species (''Cebus trinitatis'') of gracile capuchin monkey. It is found on the island of Trinidad. Taxonomy Boubli ''et al''. found in a 2012 study that th ...
, ''Cebus trinitatis'' * 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 The Santa Marta white-fronted capuchin (''Cebus malitiosus'') is a species of gracile capuchin monkey from Colombia. It was formerly considered a subspecies of the ''Cebus albifrons'' or a synonym of the Colombian white-faced capuchin (''C. capu ...
, ''Cebus malitiosus'' * Ecuadorian white-fronted capuchin, ''Cebus aequatorialis'' The placement of the
Trinidad white-fronted capuchin The Trinidad white-fronted capuchin is a subspecies (''Cebus albifrons trinitatis'') or species (''Cebus trinitatis'') of gracile capuchin monkey. It is found on the island of Trinidad. Taxonomy Boubli ''et al''. found in a 2012 study that th ...
is controversial; the
American Society of Mammalogists The American Society of Mammalogists (ASM) was founded in 1919. Its primary purpose is to encourage the study of mammals, and professions studying them. There are over 4,500 members of this society, and they are primarily professional scientists ...
classifies it as conspecific with '' C. brunneus'' based on a 2012 study later found to be flawed, while the
IUCN Red List The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is the world's most comprehensive inventory of the global conservation status of biol ...
classifies it as a distinct species (''Cebus triniatis'') due to debate over the aforementioned study, and the ITIS classifies it as a subspecies of ''C. albifrons'', also due to debate over the aforementioned study.


Taxonomic history

Philip Hershkovitz and
William Charles Osman Hill Dr William Charles Osman Hill FRSE FZS FLS FRAI (13 July 1901 – 25 January 1975) was a British anatomist, primatologist, and a leading authority on primate anatomy during the 20th century. He is best known for his nearly completed eight-vol ...
published taxonomies of the
capuchin monkey 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 fores ...
s in 1949 and 1960, respectively. These taxonomies established four species of capuchin monkey in the
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
''Cebus''. One of those species, ''
Cebus apella 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 o ...
'', is a robust capuchin and is now included in the genus ''Sapajus''. The other three ''Cebus'' species included in that taxonomy were the gracile capuchin species ''Cebus albifrons'', ''Cebus nigrivittatus'' and the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
''Cebus capucinus''. ''Cebus nigrivittatus'' was subsequently renamed ''Cebus olivaceus''. ''Cebus kaapori'' had been considered a
subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (morphology), but that can successfully interbreed. Not all species ...
of ''C. olivaceus'' but Groves (2001 and 2005) and Silva (2001) regarded it as a separate species.


Evolutionary history

The gracile capuchins, like all capuchins, are members of the family Cebidae, which also includes the squirrel monkeys. The evolution of the squirrel monkeys and capuchin monkeys is believed to have diverged about 13 million years ago. 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 The Amazon River (, ; es, Río Amazonas, pt, Rio Amazonas) in South America is the largest river by discharge volume of water in the world, and the disputed longest river system in the world in comparison to the Nile. The headwaters of t ...
, which separated the monkeys in the
Amazon Amazon most often refers to: * Amazons, a tribe of female warriors in Greek mythology * Amazon rainforest, a rainforest covering most of the Amazon basin * Amazon River, in South America * Amazon (company), an American multinational technology c ...
north of the Amazon River, which evolved into the gracile capuchins, from those in the
Atlantic Forest The Atlantic Forest ( pt, Mata Atlântica) is a South American forest that extends along the Atlantic coast of Brazil from Rio Grande do Norte state in the northeast to Rio Grande do Sul state in the south and inland as far as Paraguay and th ...
south of the river, which evolved into the robust capuchins.


Morphology

Gracile capuchins have longer limbs relative to their body size compared with robust capuchins. Gracile capuchins also have rounder skulls and other differences in skull morphology. Gracile capuchins lack certain adaptations for opening hard nuts which robust capuchins have. These include differences in the teeth and jaws, and the lack of a
sagittal crest A sagittal crest is a ridge of bone running lengthwise along the midline of the top of the skull (at the sagittal suture) of many mammalian and reptilian skulls, among others. The presence of this ridge of bone indicates that there are exceptiona ...
. Exterior differences include the fact that, although some females have tufts on their head (
Humboldt's white-fronted capuchin Humboldt's white-fronted capuchin (''Cebus albifrons'') is a species of gracile capuchin monkey. It is found in Colombia, Venezuela, Brazil, and potentially the island of Trinidad. Taxonomy The species name ''Cebus albifrons'' was formerly co ...
and
Guianan weeper capuchin The wedge-capped capuchin or Guianan weeper capuchin (''Cebus olivaceus'') 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 ...
), no male gracile capuchin has tufts, while all robust capuchins have tufts. Also, no gracile capuchins have
beard A beard is the hair that grows on the jaw, chin, upper lip, lower lip, cheeks, and neck of humans and some non-human animals. In humans, usually pubescent or adult males are able to grow beards. Throughout the course of history, societal at ...
s.


Distribution

Gracile capuchin monkeys have a wide range over
Central America Central America ( es, América Central or ) is a subregion of the Americas. Its boundaries are defined as bordering the United States to the north, Colombia to the south, the Caribbean Sea to the east, and the Pacific Ocean to the west. ...
and north and north-west
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...
. The Panamanian white-headed capuchin is the most northern species, occurring in Central America from
Honduras Honduras, officially the Republic of Honduras, is a country in Central America. The republic of Honduras is bordered to the west by Guatemala, to the southwest by El Salvador, to the southeast by Nicaragua, to the south by the Pacific Oce ...
to
Panama Panama ( , ; es, link=no, Panamá ), officially the Republic of Panama ( es, República de Panamá), is a transcontinental country spanning the southern part of North America and the northern part of South America. It is bordered by Cos ...
. The Colombian white-headed capuchin also has a northern distribution in
Colombia Colombia (, ; ), officially the Republic of Colombia, is a country in South America with insular regions in North America—near Nicaragua's Caribbean coast—as well as in the Pacific Ocean. The Colombian mainland is bordered by the Car ...
and
Ecuador Ecuador ( ; ; Quechua: ''Ikwayur''; Shuar: ''Ecuador'' or ''Ekuatur''), officially the Republic of Ecuador ( es, República del Ecuador, which literally translates as "Republic of the Equator"; Quechua: ''Ikwadur Ripuwlika''; Shuar: ''Eku ...
west of the
Andes The Andes, Andes Mountains or Andean Mountains (; ) are the longest continental mountain range in the world, forming a continuous highland along the western edge of South America. The range is long, wide (widest between 18°S – 20°S ...
. The white-fronted capuchin is found over large portions of
Colombia Colombia (, ; ), officially the Republic of Colombia, is a country in South America with insular regions in North America—near Nicaragua's Caribbean coast—as well as in the Pacific Ocean. The Colombian mainland is bordered by the Car ...
,
Peru , image_flag = Flag of Peru.svg , image_coat = Escudo nacional del Perú.svg , other_symbol = Great Seal of the State , other_symbol_type = Seal (emblem), National seal , national_motto = "Fi ...
and western
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
, as well as into southern
Venezuela Venezuela (; ), officially the Bolivarian Republic of Venezuela ( es, link=no, República Bolivariana de Venezuela), is a country on the northern coast of South America, consisting of a continental landmass and many islands and islets in th ...
and northern
Bolivia , image_flag = Bandera de Bolivia (Estado).svg , flag_alt = Horizontal tricolor (red, yellow, and green from top to bottom) with the coat of arms of Bolivia in the center , flag_alt2 = 7 × 7 square p ...
. The weeper capuchin is found over much of Venezuela and over
The Guianas The Guianas, sometimes called by the Spanish loan-word ''Guayanas'' (''Las Guayanas''), is a region in north-eastern South America which includes the following three territories: * French Guiana, an overseas department and region of France * ...
, as well as part of northern Brazil. The Kaapori capuchin has a range that is disjoint from the other gracile capuchins, living in northern Brazil within the states of
Pará Pará is a Federative units of Brazil, state of Brazil, located in northern Brazil and traversed by the lower Amazon River. It borders the Brazilian states of Amapá, Maranhão, Tocantins (state), Tocantins, Mato Grosso, Amazonas (Brazilian state) ...
and
Maranhão Maranhão () is a state in Brazil. Located in the country's Northeast Region, it has a population of about 7 million and an area of . Clockwise from north, it borders on the Atlantic Ocean for 2,243 km and the states of Piauí, Tocantins and ...
. The only species to inhabit the
Caribbean The Caribbean (, ) ( es, El Caribe; french: la Caraïbe; ht, Karayib; nl, De Caraïben) is a region of the Americas that consists of the Caribbean Sea, its islands (some surrounded by the Caribbean Sea and some bordering both the Caribbean Se ...
islands is the Trinidad white-fronted capuchin.


Behaviour


Tool use

Some gracile capuchins are known to use tools. These include
white-headed capuchin White-faced capuchin, or white headed capuchin, can refer to either of two species of gracile capuchin monkey: * ''Cebus imitator'', the Panamanian white-faced capuchin, also known as the Panamanian white-headed capuchin or Central American w ...
s rubbing secretions from leaves over their bodies, using leaves as gloves when rubbing fruit or caterpillar secretions and using tools as a probe.
White-fronted capuchin 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: Bo ...
s have been observed using leaves as a cup to drink water.


Mating systems


Male weaponry

Intrasexual selection, or male-male competition, occurs when males invoke contests in order to gain the opportunity to reproduce with a female and maximize their reproductive success. Often males are adorned with weaponry, which can be used in order to increase their chances of winning contests for possible mates. In the genus Cebus, there is a large amount of dimorphism in canine size between males and females. Canines are hypothesized to be larger in males because canine dimorphism is generally correlated to male-male competition. In the
wedge-capped capuchin The wedge-capped capuchin or Guianan weeper capuchin (''Cebus olivaceus'') 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 ove ...
there is a larger amount of canine dimorphism compared to the
white-faced capuchin White-faced capuchin, or white headed capuchin, can refer to either of two species of gracile capuchin monkey: * ''Cebus imitator'', the Panamanian white-faced capuchin, also known as the Panamanian white-headed capuchin or Central American whi ...
and the
white-fronted capuchin 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: Bo ...
. The difference in canine dimorphism between these species can be correlated to the differences in social structure of these three groups. The alpha male of the wedge-capped capuchin tends to monopolize mating, therefore engaging in more male-male competition, while in the white-faced capuchin and in the white-fronted capuchin the alpha male does not monopolize mating and allows subordinate males to mate with females. While not much is known about the
Kaapori capuchin The Kaapori capuchin (''Cebus kaapori''), 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 populat ...
, due to its low population size, it is likely it would possess more canine dimorphism, like the wedge-capped capuchin, because of its similar social structure with a monopolizing alpha male and peripheral subordinate males.


Direct and indirect female benefits

If a female is presented an opportunity to copulate with a male she will evaluate both the costs and benefits of that male. Females can obtain direct benefits from males she mates with, where the female gains an instant benefit from the male to herself. Direct benefits that would apply to females of the genus ''Cebus'' would include; vigilance from males, protection from predators and conspecifics, and increased resources. Females can also benefit indirectly from males, in the form of phenotypic and genotypic benefits to her offspring as well as male protection of those offspring. Alpha males are more fit, and therefore more likely, to provide direct and indirect benefits to the female compared to other subordinate males. In the white-faced capuchin the alpha male fathers 70-90% of the offspring produced by females in his group. It is hypothesized that females are mating with alpha males while they are ovulating and then mating with subordinate males after they are no longer conceptive. Some female primates, like in the white-fronted capuchin, will mate will subordinate males while they are no longer conceptive in order to decrease the amount of resource competition and increase the amount of male protection for her offspring.


Parental care

Capuchin infants are born in an
altricial In biology, altricial species are those in which the young are underdeveloped at the time of birth, but with the aid of their parents mature after birth. Precocial species are those in which the young are relatively mature and mobile from the mome ...
state, which means they need a lot of parental care in order to survive. The majority of parental care in the genus ''Cebus'' is provided by the mother, but in the case of the wedge-capped capuchin, parental care is also provided by other conspecific females; this type of care is referred to as allomaternal care. In the wedge-capped capuchin, the mother will provide the infant care for the first three months, however for the next three months the infant relies on the care of other females. In agreement with kin selection theory, kin of the mother are more likely to provide care to the infant compared to other females in the group; siblings were four times as likely to provide infant care compared to other group females. Male parental care is rare in the genus ''Cebus'', only in the white-headed capuchin is there some interaction between males and offspring. In white-headed capuchins males will often investigate, or at least tolerate, their offspring. Alpha males are also more likely to interact with their offspring than subordinate males.


Conservation status

All gracile capuchin species except the Kaapori capuchin are rated as least concern by the
International Union for Conservation of Nature The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
. The Kaapori capuchin is rated as critically endangered.


References

{{Taxonbar, from=Q8447051 ^