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The black howler (''Alouatta caraya'') or black-and-gold howler, is among the largest
New World monkey New World monkeys are the five families of primates that are found in the tropical regions of Mexico, Central and South America: Callitrichidae, Cebidae, Aotidae, Pitheciidae, and Atelidae. The five families are ranked together as the Ceboi ...
s and a member of the ''
Alouatta Howler monkeys (genus ''Alouatta'', monotypic in subfamily Alouattinae) are the most widespread primate genus in the Neotropics and are among the largest of the platyrrhines along with the muriquis (''Brachyteles''), the spider monkeys (''Atele ...
''
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
. The black howler is distributed in areas of South America such as
Paraguay Paraguay, officially the Republic of Paraguay, is a landlocked country in South America. It is bordered by Argentina to the Argentina–Paraguay border, south and southwest, Brazil to the Brazil–Paraguay border, east and northeast, and Boli ...
, southern
Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is the world's List of countries and dependencies by area, fifth-largest country by area and the List of countries and dependencies by population ...
, eastern
Bolivia Bolivia, officially the Plurinational State of Bolivia, is a landlocked country located in central South America. The country features diverse geography, including vast Amazonian plains, tropical lowlands, mountains, the Gran Chaco Province, w ...
, northern
Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...
, and
Uruguay Uruguay, officially the Oriental Republic of Uruguay, is a country in South America. It shares borders with Argentina to its west and southwest and Brazil to its north and northeast, while bordering the Río de la Plata to the south and the A ...
. This species is
sexually dimorphic Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
, with adult males having entirely black fur and adult females and babies of both sexes having an overall golden colouring; which emphasizes black-and-gold in the name. The
IUCN The International Union for Conservation of Nature (IUCN) is an international organization working in the field of nature conservation and sustainable use of natural resources. Founded in 1948, IUCN has become the global authority on the status ...
Red List has classed the black howler as
Near Threatened A near-threatened species is a species which has been Conservation status, categorized as "Near Threatened" (NT) by the International Union for Conservation of Nature (IUCN) as that may be vulnerable to Endangered species, endangerment in the ne ...
as a result of a recent population reduction due to a variety of human-caused factors.


Taxonomy

The black howler is one of nine species of the ''
Alouatta Howler monkeys (genus ''Alouatta'', monotypic in subfamily Alouattinae) are the most widespread primate genus in the Neotropics and are among the largest of the platyrrhines along with the muriquis (''Brachyteles''), the spider monkeys (''Atele ...
'' genus that have been identified and is in the
Atelidae The Atelidae are one of the five families of New World monkeys now recognised. It was formerly included in the family Cebidae. Atelids are generally larger monkeys; the family includes the howler, spider, woolly, and woolly spider monkeys (t ...
family. Because of this species' sexual dimorphism, the taxon for the black howler has been distinguished. * ''Alouatta caraya'' * '' Alouatta pigra'' * ''Alouatta seniculus'' * ''Alouatta arctoidea'' * ''Alouatta sara'' * ''Alouatta macconnelli'' * ''Alouatta guariba'' * ''Alouatta belzebul'' * '' Alouatta palliata''


Description


Physical appearance

Black howlers are
sexually dimorphic Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
, the male and the female are distinguishable due to the difference in their fur coatings. As the male black howler grows older, his fur will change from a golden colour to a complete black fur coat. The female, as well as infants of both sexes, have fur of a golden tone. Female howlers are lighter in weight than males, who are 47% heavier. Males' weight ranges from 6 kg to 7.7 kg. The
hyoid bone The hyoid-bone (lingual-bone or tongue-bone) () is a horseshoe-shaped bone situated in the anterior midline of the neck between the chin and the thyroid-cartilage. At rest, it lies between the base of the mandible and the third cervical verte ...
volume of male black howlers is larger than that of female black howlers; it is assumed that this bone serves as a chamber for their howls. They have
prehensile tail A prehensile tail is the tail of an animal that has Adaptation (biology), adapted to grasp or hold objects. Fully Prehensility, prehensile tails can be used to hold and manipulate objects, and in particular to aid arboreal creatures in finding and ...
s with a bare patch of skin on the underside, making them versatile climbers and allowing them to comfortably stay around in the canopy of trees.


Behaviour and diet

Black howlers share many behaviors with other
howler monkey Howler monkeys (genus ''Alouatta'', monotypic in subfamily Alouattinae) are the most widespread primate genus in the Neotropical realm, Neotropics and are among the largest of the New World monkey, platyrrhines along with the muriquis (''Brachyte ...
s. They are recognized for their 'howl', a powerful vocalization. A howler's 'howl' is one of the primates' strongest vocalizations, and it may be heard up to two kilometers away. The vocalizations of black howlers have been studied in Brazil, where they mostly vocalize around dawn. Howling is used as a behavioural strategy for displaying territory and regulating inter-group distance. This species is mostly arboreal, rarely coming down to the ground. They are also one of the least active groups of primates, as they are diurnal and spend up to 60-70% of the day resting or sleeping. Although the diet of the black howler is largely folivorous, it does not prevents the digestion of fruits, figs, mosses, bark, seeds, and flowers. Eating leaves may appear to be unfeasible for other primates, as it lacks nourishment and energy; however, howlers have developed a complicated digestive system that allows them to digest leaves quickly and have strong molar teeth to help chew mature leaves. Black howlers' activity levels vary depending on their diet. During the winter months, when fruit is scarce, most of their diet consists of leaves and they are less active compared to the summer months when fruit is more available. Activity ranges between feeding, socialization, locomotion, and sexual activity. Their gut
microbiome A microbiome () is the community of microorganisms that can usually be found living together in any given habitat. It was defined more precisely in 1988 by Whipps ''et al.'' as "a characteristic microbial community occupying a reasonably wel ...
composition changes seasonally to accommodate dietary changes due to restrictions of resources in the environment. Their diet is not different between different sexes and ages.


Group composition

Group composition has a large varied range in population density and social organization when focusing on black howler monkeys. Due to minimal human influence along the
Paraná River The Paraná River ( ; ; ) is a river in south-central South America, running through Brazil, Paraguay, and Argentina for some ."Parana River". Encyclopædia Britannica. Encyclopædia Britannica Online. Encyclopædia Britannica Inc., 2012. Web. ...
in
Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is the world's List of countries and dependencies by area, fifth-largest country by area and the List of countries and dependencies by population ...
, for example, the woods remain well conserved, which results in larger densities of black howlers. Usually, howler monkeys have a single dominant male, with multiple adult females and babies in smaller groups. In larger groups, the howler monkeys' group composition can vary. In a 2009 study by Lucas M. Aguiar, Gabriela Ludwig, and Fernando C. Passos, in the Upper Paraná River of southern Brazil, the black and gold howler groups size was around 6 to 18 individuals. There were 11 groups total in Paraná River and groups were all multi-male; the females were outnumbered. The range for how many black howler monkeys in terms of age and sex in individual groups were represented. The study concluded that in these individual groups, it was likely to see a range of male adult black howlers from two to five, three to seven adult females, zero to four adolescents, and zero to three babies.


Mating

Mating behaviour and patterns of the black howlers have been identified. A study by Martin M. Kawoleski and Paul A. Garber in 2010 examined the promiscuity of black howler females along the
Paraná River The Paraná River ( ; ; ) is a river in south-central South America, running through Brazil, Paraguay, and Argentina for some ."Parana River". Encyclopædia Britannica. Encyclopædia Britannica Online. Encyclopædia Britannica Inc., 2012. Web. ...
in
Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...
using a scan sampling technique. According to the findings, sexual solicitation of female
mate choice Mate choice is one of the primary mechanisms under which evolution can occur. It is characterized by a "selective response by animals to particular stimuli" which can be observed as behavior.Bateson, Paul Patrick Gordon. "Mate Choice." Mate Choi ...
occurred when a female actively pursued a particular male by grooming, touching, and following him, and displaying a specific posture to get the male to approach them. During fertile and nonfertile phases, black howler monkey species' females mate with many adult males. Female black and gold howlers have been shown to conceive within a 3–4 day ovulatory phase, according to previous research. It is still unknown if male howlers can discriminate ovulatory phases in females using visual or smell cues. The adult female black howlers tend to give birth within 180–194 days after copulating. In black howlers, mating promiscuity is a behavioural habit that favours offspring genetic variety and can lead to a lot of genetic variations inside the group. According to Kawoleski and Garber, the black howler has a low rate of
infanticide Infanticide (or infant homicide) is the intentional killing of infants or offspring. Infanticide was a widespread practice throughout human history that was mainly used to dispose of unwanted children, its main purpose being the prevention of re ...
, which is one of the theories said to contribute to the female howler's promiscuity.


Distribution and habitat

The black howler is distributed in areas of Central and South America such as
Paraguay Paraguay, officially the Republic of Paraguay, is a landlocked country in South America. It is bordered by Argentina to the Argentina–Paraguay border, south and southwest, Brazil to the Brazil–Paraguay border, east and northeast, and Boli ...
, southern
Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is the world's List of countries and dependencies by area, fifth-largest country by area and the List of countries and dependencies by population ...
, eastern
Bolivia Bolivia, officially the Plurinational State of Bolivia, is a landlocked country located in central South America. The country features diverse geography, including vast Amazonian plains, tropical lowlands, mountains, the Gran Chaco Province, w ...
, northern
Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...
, and occasionally,
Uruguay Uruguay, officially the Oriental Republic of Uruguay, is a country in South America. It shares borders with Argentina to its west and southwest and Brazil to its north and northeast, while bordering the Río de la Plata to the south and the A ...
. Howler monkeys are adaptable to numerous forest environments in Central and South America. They have been spotted in tropical and subtropical forests such as evergreen, seasonally dry deciduous, semideciduous, gallery, montane, woodland, and flooded forests. They may survive in a variety of settings, including disturbed ecosystems and forest remnants in agricultural regions and human populations.


Status and conservation

From 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 an inventory of the global conservation status and extinction risk of biological ...
, the black howler is seen as a
Near Threatened A near-threatened species is a species which has been Conservation status, categorized as "Near Threatened" (NT) by the International Union for Conservation of Nature (IUCN) as that may be vulnerable to Endangered species, endangerment in the ne ...
species. This is a result to population declines, habitat degradation, hunting pressure, and disease outbreak vulnerability. Anthropogenic disturbances are the top common reasons for the slow decrease of the black howler population; jeopardizing and degrading their ecosystems for residential and commercial development as well as agriculture. Hunting and the illegal pet trade are also contributing factors to the black howler's population reduction. In a 2020 study in the city of Pilar,
Paraguay Paraguay, officially the Republic of Paraguay, is a landlocked country in South America. It is bordered by Argentina to the Argentina–Paraguay border, south and southwest, Brazil to the Brazil–Paraguay border, east and northeast, and Boli ...
, examining the attitudes of humans towards urban howler monkeys, the researchers asked selected participants questions regarding hunting and pet trade of black and gold howler monkeys. In their findings, 93% of participants said they knew it was illegal to kill the monkeys, but more than half of the participants did not know it was also prohibited to keep them as pets. It was also stated in their findings that 8% of participants believe that black and gold howler monkeys had been hunted for their flesh. Black howlers are also susceptible to disease such as yellow fever. In Brazil, there have been outbreaks of sylvatic yellow fever that have resulted in significant population declines.


Human interactions

The interaction between people and black and gold howlers was also a topic of the research conducted by Marco Alesci et al. in 2020 in the city of
Pilar, Paraguay Pilar () is the capital city of the Paraguayan Departments of Paraguay, department of Ñeembucú, located along the Paraguay River in the southwestern part of the country. Located about from Asunción, Pilar serves as an important center of com ...
. People interviewed for this research believed that there were many benefits to having the black howler monkeys in the urban environment such as a benefit for increasing tourism. The researchers also polled a group of people in Pilar regarding the conservation of black howlers in the area as part of their research. As a result, domestic dogs attacking black howlers in urban areas, electrocution on uninsulated power lines, and people killing the monkeys were all recognized as threats. Because the power lines are commonly utilized for transit by the black howler monkeys, three howler fatalities by electrocution were documented during the research period. As mentioned in the article, efforts for the conservation of the black howlers in Pilar are minimal, but if urbanization keeps rising, it may pose serious dangers to their existence by diminishing accessible trees and pushing howlers to walk on power wires or on the ground more.


References

{{Authority control black howler Mammals of Argentina Primates of Brazil Mammals of Bolivia Mammals of Paraguay black howler Taxa named by Alexander von Humboldt