Howler monkeys [1] | |
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Brown howler monkey | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Primates |
Suborder: | Haplorhini |
Infraorder: | Simiiformes |
Family: | Atelidae |
Subfamily: | Alouattinae Trouessart, 1897 (1825) |
Genus: | Alouatta Lacépède, 1799 |
Type species | |
Simia belzebul Linnaeus, 1766 | |
Species | |
See text | |
Alouatta distribution | |
Synonyms | |
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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 (Ateles) and woolly monkeys (Lagotrix). The monkeys are native to South and Central American forests. They are famous for their loud howls, which can be heard up to three miles away through dense rain forest. [2] Fifteen species are recognized. Previously classified in the family Cebidae, they are now placed in the family Atelidae. They are primarily folivores but also significant frugivores, acting as seed dispersal agents through their digestive system and their locomotion. Threats include human predation, habitat destruction, illegal wildlife trade, and capture for pets or zoo animals.
Group | Image | Scientific name | Subspecies | Distribution |
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A. palliata group | Coiba Island howler, Alouatta coibensis |
| Panama | |
Mantled howler, Alouatta palliata |
| Colombia, Costa Rica, Ecuador, Guatemala, Honduras, Mexico, Nicaragua, Panama and Peru | ||
Guatemalan black howler, Alouatta pigra | Belize, Guatemala and Mexico | |||
A. seniculus group | Ursine howler, Alouatta arctoidea | Venezuela and possibly Colombia | ||
Red-handed howler, Alouatta belzebul | Brazil | |||
Spix's red-handed howler, Alouatta discolor | Brazil | |||
Brown howler, Alouatta guariba |
| southeastern Brazil and far northeastern Argentina (Misiones) | ||
Juruá red howler, Alouatta juara | Peru and Brazil | |||
Guyanan red howler, Alouatta macconnelli | Suriname, Guyana, Trinidad, French Guiana, Venezuela and Brazil | |||
Amazon black howler, Alouatta nigerrima | Brazil | |||
Purus red howler, Alouatta puruensis | Brazil, Peru and north of Bolivia | |||
Bolivian red howler, Alouatta sara | Bolivia | |||
Venezuelan red howler, Alouatta seniculus | Venezuela, Colombia, Ecuador, Peru and Brazil | |||
Maranhão red-handed howler, Alouatta ululata | Brazil | |||
A. caraya group | Black howler, Alouatta caraya | Paraguay, southern Brazil, eastern Bolivia, northern Argentina, and Uruguay | ||
Howler monkeys have short snouts and wide-set, round nostrils. Their noses are very keen, and they can smell out food (primarily fruit and nuts) up to 2 km away. Their noses are usually roundish snout-type, and the nostrils have many sensory hairs growing from the interior. They range in size from 56 to 92 cm (22 to 36 in), excluding their tails, which can be equally long; in fact in some cases the tail has been found to be almost five times the body length.[ citation needed ] This is a prime characteristic. Like many New World monkeys, they have prehensile tails, which they use while picking fruit and nuts from trees. Unlike other New World monkeys, both male and female howler monkeys have trichromatic color vision. [3] This has evolved independently from other New World monkeys due to gene duplication. [4] They have lifespans of 15 to 20 years. Howler species are dimorphic and can also be dichromatic (i.e. Alouatta caraya). Males are typically 1.5 to 2.0 kg heavier than females.
Males experience an evolutionary trade off between investments in precopulatory traits, larger hyoids but smaller testes, or post-copulatory traits, larger testes and smaller hyoids.The hyoid of Alouatta is pneumatized, one of the few cases of postcranial pneumaticity outside the Saurischia. The volume of the hyoid of male howler monkeys is negatively correlated with the dimensions of their testes, and with the number of males per group. Larger hyoids decrease space between formant, and offers impression of larger body size. [5] They have a flat cranial shape due to a folivorous diet and an advanced vocal system. Their brain growth is posterior rather than superior or inferior as in other platyrrhines. [6]
Howler monkeys generally move quadrupedally on the tops of branches, usually grasping a branch with at least two hands or one hand and the tail at all times. Their strong prehensile tails are able to support their entire body weight. Fully grown adult howler monkeys do not often rely on their tails for full-body support, but juveniles do so more frequently. A significant amount of their travel is done through the ground, with sitting and resting being their most frequent postures. [7]
Most howler species live in groups of six to 15 animals, with one to three adult males and multiple females. Mantled howler monkeys are an exception, commonly living in groups of 15 to 20 individuals with more than three adult males. The number of males in a given group is inversely correlated with the size of their hyoids and is positively correlated with testes size.
This results in two distinct groups, wherein one male with a larger hyoid and smaller testes copulates exclusively with a group of females, suggesting precopulatory vocal competition. The other group has more males, which have smaller hyoids, and larger testes. The larger the number of males, the smaller the hyoid, and the larger the testes. [5] Female howler monkeys breed with multiple males within their group, with males in neighboring groups, and with solitary males. Central males tie up fellowship with cycling females. [8] Unlike most New World monkeys, in which one sex remains in natal groups, juveniles of both sexes emigrate from their natal groups, [9] such that howler monkeys could spend the majority of their adult lives in association with unrelated monkeys.
Physical fighting among group members is infrequent and generally of short duration, but serious injuries can result. Both males and females rarely fight with each other, but physical aggression is even more rare between sexes. [9] [10] Group size varies by species and by location, with an approximate ratio of one male to four females. [9]
As their name suggests, vocal communication forms an important part of their social behavior. They each have an enlarged basihyal or hyoid bone, which helps them make their loud vocalizations. Group males generally call at dawn and dusk, as well as interspersed times throughout the day. Their main vocals consist of loud, deep, guttural growls or "howls". Howler monkeys are widely considered to be the loudest land animals. According to Guinness Book of World Records, their vocalizations can be heard clearly for 3 mi (4.8 km). [11] The function of howling is thought to relate to intergroup spacing and territory protection, as well as possibly to mate-guarding. Howlers call usually when they are in areas with major feeding sites, which in some sort lead to advertise major feeding sites and their willingness to defend locally available fruit trees. Black howler monkeys incorporate information on resource availability along with neighbors’ current location. And the abundance of flowers are found to be an important factor that influenced behavior. Neighbors are more likely to move towards these calls when resource are scarce, and the reverse is true. [12]
These large and slow-moving monkeys are the only folivores of the New World monkeys. Howlers eat mainly top canopy leaves, together with fruit, buds, flowers, and nuts. They need to be careful not to eat too many leaves of certain species in one sitting, as some contain toxins that can poison them. [13] Howler monkeys are also known to occasionally raid birds' nests, chicken coops, and consume the eggs. [14] When in smaller groups (up to twelve individuals) and low rainfall (up to 2,200 mm (87 in)), they are more frugivorous. In larger groups and increased rainfall, frugivory decreases as a result of competition and fast food depletion. [15] As they digest fruit, more than 90% of the fruits' seeds are excreted without damage, which results in seed dispersal and distribution in tropical forests. [16]
Howlers use the upper-middle part of their sleeping tree and use large branches on the 70% of nights that potentially allow for grouped sleeping or resistance to weather conditions and risk of branch breaking. Their sleeping sites are usually close to morning feeding sites. [17]
While they are not usually aggressive, brown howler monkeys do not take well to captivity and are of bad-tempered and unfriendly disposition. However, the black howler monkey (Alouatta caraya) is a relatively common pet in contemporary Argentina due to its gentle nature (in comparison to the capuchin monkey's aggressive tendencies), in spite of its lesser intelligence, as well as the liabilities of the size of its droppings and the male monkey’s loud vocalizations.
John Lloyd Stephens described the howler monkeys at the Maya ruins of Copán as "grave and solemn, almost emotionally wounded, as if officiating as the guardians of consecrated ground". To the Mayas of the Classic period, they were the divine patrons of the artisans, especially scribes and sculptors. They were seen as gods in some tribes, and the long, sleek tail was worshipped for its beauty. Copán, in particular, is famous for its representations of howler monkey gods. Two howler monkey brothers play a role in the myth of the Maya Hero Twins included in the Popol Vuh , a widely feared tale of soul and passion.
Primates is an order of mammals, which is further divided into the strepsirrhines, which include lemurs, galagos, and lorisids; and the haplorhines, which include tarsiers and simians. Primates arose 85–55 million years ago first from small terrestrial mammals, which adapted for life in tropical forests: many primate characteristics represent adaptations to the challenging environment among tree tops, including large brain sizes, binocular vision, color vision, vocalizations, shoulder girdles allowing a large degree of movement in the upper limbs, and opposable thumbs that enable better grasping and dexterity. Primates range in size from Madame Berthe's mouse lemur, which weighs 30 g (1 oz), to the eastern gorilla, weighing over 200 kg (440 lb). There are 376–524 species of living primates, depending on which classification is used. New primate species continue to be discovered: over 25 species were described in the 2000s, 36 in the 2010s, and six in the 2020s.
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 Ceboidea, the only extant superfamily in the parvorder Platyrrhini.
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. They are found throughout the forested regions of Central and South America, from Mexico to northern Argentina.
The Colombian red howler or Venezuelan red howler is a South American species of howler monkey, a type of New World monkey, found in the western Amazon Basin in Venezuela, Colombia, Ecuador, Peru and Brazil. The population in the Santa Cruz Department in Bolivia was split off as a separate species, the Bolivian red howler, in 1986, and more recently, splitting off the population in northeastern South America and Trinidad as the Guyanan red howler has occurred. All howler monkeys belong to the family Atelidae and the infraorder Platyrrhini.
The brown howler, also known as brown howler monkey, is a species of howler monkey, a type of New World monkey that lives in forests in south-eastern Brazil and far north-eastern Argentina (Misiones). It lives in groups of two to 11 individuals. Despite the name "brown howler", it is notably variable in colour, with some individuals appearing largely reddish-orange or black.
The black howler or black-and-gold howler, is among the largest New World monkeys and a member of the Alouatta genus. The black howler is distributed in areas of South America such as Paraguay, southern Brazil, eastern Bolivia, northern Argentina, and Uruguay. This species is sexually dimorphic, 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 Red List has classed the black howler as Near Threatened as a result of a recent population reduction due to a variety of human-caused factors.
The red-handed howler is a vulnerable species of howler monkey, a type of New World monkey. It is endemic to Brazil, found in the southeastern Amazon and disjunctly in the Atlantic Forest between Rio Grande do Norte and Sergipe.
The white-bellied spider monkey, also known as the white-fronted or long-haired spider monkey, is an endangered species of spider monkey, a type of New World monkey. It is found in the north-western Amazon in Colombia, Ecuador, Venezuela, Peru and Brazil, ranging as far south as the lower Ucayali River and as far east as the Branco River. In the past, the Peruvian, brown and white-cheeked spider monkeys have been treated as subspecies of A. belzebuth. As presently defined, the white-bellied spider monkey is monotypic. It has a whitish belly and a pale patch on the forehead, which, despite its common name, often is orange-buff, though this might be due to dirt and other stainers. They live in groups of 20 to 40 individuals, splitting into small parties of 1 to 9 when in activity.
The Bolivian red howler is a species of howler monkey, a type of New World monkey, endemic to Bolivia. It can be found in rain forests, including riverine and seasonally flooded forests.
The mantled howler is a species of howler monkey, a type of New World monkey, from Central and South America. It is one of the monkey species most often seen and heard in the wild in Central America. It takes its "mantled" name from the long guard hairs on its sides.
The Yucatán black howler, or Guatemalan black howler, is a species of howler monkey, a type of New World monkey, from Central America. It is found in Belize, Guatemala and Mexico, in and near the Yucatán Peninsula. It lives in evergreen, semideciduous and lowland rain forests. It is also known as the baboon in Belize, although it is not closely related to the baboons in Africa.
The Coiba Island howler is a type of howler monkey, a type of New World monkey, endemic to Panama. Although the Coiba Island howler has been recognized as a separate species by a number of authorities since a 1987 study of its fingerprints, mitochondrial DNA testing found it does not differ from mantled howler populations in any significant way. A reason given for treating it as a separate species is that the dermal ridges of its hands and feet differ from those of the mantled howler.
The evolution of color vision in primates is highly unusual compared to most eutherian mammals. A remote vertebrate ancestor of primates possessed tetrachromacy, but nocturnal, warm-blooded, mammalian ancestors lost two of four cones in the retina at the time of dinosaurs. Most teleost fish, reptiles and birds are therefore tetrachromatic while most mammals are strictly dichromats, the exceptions being some primates and marsupials, who are trichromats, and many marine mammals, who are monochromats.
Sexual dimorphism describes the morphological, physiological, and behavioral differences between males and females of the same species. Most primates are sexually dimorphic for different biological characteristics, such as body size, canine tooth size, craniofacial structure, skeletal dimensions, pelage color and markings, and vocalization. However, such sex differences are primarily limited to the anthropoid primates; most of the strepsirrhine primates and tarsiers are monomorphic.
The Mexican howler is a subspecies of the mantled howler, A. palliata. This subspecies is found predominantly in forests between south eastern Mexico and north eastern Peru. Typical of its species, the Mexican howler monkey has a prehensile tail, a deep jaw, and a large pharynx which it uses to make characteristically deep and resonating howls. Mantled howler monkeys are known for forming unusually large cohorts averaging 14 members and sometimes extending to 40 members.
The ursine howler is a species of howler monkey native to Venezuela and possibly Colombia. It is sometimes considered a subspecies of the Venezuelan red howler and classified as Alouatta seniculus arctoidea.
Spix's red-handed howler is a species of Howler monkey native to the southeastern Amazon in Brazil. It is threatened by deforestation and hunting.
The southern brown howler is a monkey subspecies of brown howler native to southeastern Brazil and far northeastern Argentina (Misiones). Gregorin, 2006, considered the southern brown howler to be a separate species, Alouatta clamitans, but this has not been universally accepted.
The northern brown howler is the type subspecies of the brown howler, native to Brazil. It is listed as critically endangered, with fewer than 250 individuals restricted to the vicinity of the Jequitinhonha River. The species feeds on fruits, flowers, and by preference immature leaves which are easier to digest than mature leaves; foraging for these foods in hillside habitats was shown to require more energy expenditure than in valley habitats.