Howler monkey

howler monkey[1]
Black Howler Alouatta caraya
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Primates
Family:	Atelidae
Subfamily:	Alouattinae Trouessart, 1897 (1825)
Genus:	Alouatta Lacepede, 1799
Type species
Simia belzebul Linnaeus, 1766
Species
Alouatta arctoidea Alouatta belzebul Alouatta caraya Alouatta coibensis Alouatta discolor Alouatta guariba Alouatta juara Alouatta macconnelli Alouatta nigerrima Alouatta palliata Alouatta pigra Alouatta puruensis Alouatta sara Alouatta seniculus Alouatta ululata

Howler monkeys (genus Alouatta monotypic in subfamily Alouattinae) are among the largest of the New World monkeys. Fifteen species are currently recognised. Previously classified in the family Cebidae, they are now placed in the family Atelidae. These monkeys are native to South and Central American forests. Threats to howler monkeys include human predation, habitat destruction and being captured for captivity as pets or zoo animals.

Classification

• Family Atelidae
  • Subfamily Niggroma
    • A. palliata group
      • Coiba Island Howler, Alouatta coibensis
        • Alouatta coibensis coibensis
        • Azuero Howler, Alouatta coibensis trabeata
      • Mantled Howler, Alouatta palliata
        • Ecuadorian Mantled Howler, Alouatta palliata aequatorialis
        • Golden-mantled Howler, Alouatta palliata palliata
        • Mexican Howler Monkey, Alouatta palliata mexicana
      • Guatemalan Black Howler, Alouatta pigra
    • A. seniculus group
      • Ursine Howler, Alouatta arctoidea
      • Red-handed Howler, Alouatta belzebul
      • Spix's Red-handed Howler, Alouatta discolor
      • Brown Howler, Alouatta guariba
        • Northern Brown Howler, Alouatta guariba guariba
        • Southern Brown Howler, Alouatta guariba clamitans
      • Juruá Red Howler, Alouatta juara
      • Guyanan Red Howler, Alouatta macconnelli
      • Amazon Black Howler, Alouatta nigerrima
      • Purus Red Howler, Alouatta puruensis
      • Bolivian Red Howler, Alouatta sara
      • Venezuelan Red Howler, Alouatta seniculus
      • Maranhão Red-handed Howler, Alouatta ululata
    • A. caraya group
      • Black Howler, Alouatta caraya
  • Subfamily Atelinae

Anatomy and physiology

Male Mantled Howler, Costa Rica

Howler monkeys have a short snout, and wide-set, round nostrils. They range in size from 56 to 92 cm, excluding their tail which can be equally as long. Like many New World monkeys, they have prehensile tails. Unlike other New World monkeys, both male and female howler monkeys have trichromatic color vision.^[2] This has evolved independently from other New World monkeys due to gene duplication.^[3] They have a lifespan of 15 to 20 years. Howler species are dimorphic and can also be dichromatic (i.e. Alouatta caraya). Males are, on average, 1.5 to 2 kg heavier than females.

The hyoid of Alouatta is pneumatized -- one of the few cases of postcranial pneumaticity outside Saurischia.

Locomotion

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. They have strong prehensile tails which are able to support the monkey's entire body weight. However, fully adult howler monkeys do not often rely on their tail for full body support whereas juveniles do so more frequently.

Behavior

Social systems

Most howler monkey species live in groups of 10 to 15 animals, with one to three adults males and multiple females. Mantled howlers monkeys (Alouatta palliata) are an exception, commonly living in groups of 15 to 20 individuals with more than three adult males. Unlike most New World monkeys, in which one sex remains in natal groups, juveniles of both sexes emigrate from their natal groups,^[4] such that howler monkeys could spend the majority of their adult lives in association with non-kin.

Physical fighting among group members is infrequent and generally of short duration. However, serious injuries can result. Both males and females may fight with each other but physical aggression is even more rare between sexes.^[4][5] Group size varies by species and by location, with an approximate male to female ratio of a male to four females.^[4]

Communication

A pair of howler monkeys vocalising

As their name suggests, vocal communication forms an important part of their social behavior. They 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. The main vocals consist of loud, deep guttural growls or "howls." Howler monkeys are widely considered to be the loudest land animal. According to Guinness Book of World Records, their vocalizations can be heard clearly for 20 miles (32 km). It is hypothesized that the function of howling relates to intergroup spacing and territory protection, as well as possibly mate-guarding.

Diet and feeding

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 much of certain species of mature leaf in one sitting, as some of the leaves they eat contain toxins that can poison the monkey.^[6] Howler monkeys are also known to occasionally raid birds nests and chicken coops and consume the eggs.^[7]

Relationship with humans

Howler monkey

While seldom aggressive, howler monkeys do not take well to captivity and are of surly disposition, and hence are the only monkey in their forests not made a pet by the Native Americans^{[citation needed]}. However, the Black Howler (Alouatta caraya) is a relatively common pet monkey 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 meant by the size of its droppings and the males' loud vocalisation.

Alexander von Humboldt said about howler monkeys that "their eyes, voice, and gait are indicative of melancholy", while John Lloyd Stephens described those at the Maya ruins of Copán as "grave and solemn 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. Copán in particular is famous for its representations of Howler Monkey Gods. Two howler monkey brothers play a role in the 16th century myth of the Maya Hero Twins included in the Popol Vuh.

References

1. ^ Groves, C. (2005). Wilson, D. E., & Reeder, D. M, eds. ed. Mammal Species of the World (3rd ed.). Baltimore: Johns Hopkins University Press. pp. 148–152. OCLC 62265494. ISBN 0-801-88221-4. http://www.bucknell.edu/msw3. 
2. ^ Jacobs, G. H.; Neitz, M., Deegan, J. F., & Neitz, J. (1996). "Trichromatic colour vision in New World monkeys". Nature 382 (6587): 156–158. doi:10.1038/382156a0. PMID 8700203. 
3. ^ Lucas, P. W., and N. J. Dominy (2003). "Evolution and function of routine trichromatic vision in primates". Evolution 57 (11): 2636–43. doi:10.1554/03-168. PMID 14686538. 
4. ^ ^{a b c} Sussman, R. (July 2003). Primate Ecology and Social Structure, Vol. 2: New World Monkeys, Revised First Edition. Pearson Prentice Hall. pp. 142–145. ISBN 0536743649. 
5. ^ Crockett (1997-10-02). "Family Feuds". In Ciochon, R. L., Nisbett, R. A.. Primate Anthology, The: Essays on Primate Behavior, Ecology and Conservation from Natural History. Prentice Hall. pp. 32. ISBN 9780136138457. 
6. ^ Glander, Kenneth E. (March 1977). "Poison in a monkey's Garden of Eden". Natural history 86: 146–151. 
7. ^ [1], additional text.

External links

• Primate Info Net Alouatta Factsheets
• Howler monkey audio recording
• Information about howler monkeys from Belize Zoo (photos, video and audio included)
• "The Loudest Animal in the New World," Smithsonian National Zoological Park