Avahi occidentaliswestern woolly lemur

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Geographic Range

Western woolly lemurs are found only in a restricted region in Madagascar. They are found north and east of the Betsiboka river from the Ankarafantsika Reserve to the Bay of Narinda. In addition, enclaves can be found further north on the island, in the Manongarivo Special Reserve. (Harcourt and Thornback, 1990; Mittermeier, et al., 2006)

Habitat

Western woolly lemurs are found in both tropical dry deciduous and secondary forests, occasionally reaching high population densities. They are typically observed in the forest canopy. (Harcourt and Thornback, 1990; Mittermeier, et al., 2006; Warren, 1997)

Physical Description

Western woolly lemur head-body length ranges from 25 to 29 centimeters, with the tail significantly longer at 31 to 37 centimeters. Total length ranges from 56 to 65 centimeters. This makes western woolly lemurs the smallest species of Indriidae. The dorsal fur is dense and tightly curled, hence the common name "woolly lemurs." Body coloration is light or medium grey, with some individuals exhibiting brown or olive flecks. The fur lightens in color as it approaches the rear of the animal. The tail is normally the same shade of grey as the body, but some animals have been noted to have red tails. Fur on the face, throat, and cheeks is white, distinguishing the species from Avahi laniger, which has brown fur in these areas. (Mittermeier, et al., 2006; Thalmann and Geissmann, 2000; Thalmann, 2001)

  • Sexual Dimorphism
  • sexes alike
  • Range mass
    700 to 900 g
    24.67 to 31.72 oz
  • Range length
    56 to 65 cm
    22.05 to 25.59 in

Reproduction

Western woolly lemurs are monogamous. Breeding pairs live together with their offspring year-round. (Garbutt, 2007; Harcourt and Thornback, 1990)

Like all other indriids, western woolly lemurs give birth to one offspring at a time. Breeding is seasonal, occurring from April to May, with birthing following 4 to 5 months later in August and September after a gestation period of 120 to 150 days. (Fleagle, 1999; Ganzhorn, et al., 1985; Harcourt and Thornback, 1990; Klopfer and Boskoff, 1979)

  • Breeding interval
    Western woolly lemurs breed once yearly.
  • Breeding season
    Western woolly lemurs breed from April to May.
  • Average number of offspring
    1
  • Range gestation period
    120 to 150 days
  • Average time to independence
    2 years

Mothers initially carry infants on the ventral side, or belly, then later switch infants to their dorsal side. Young woolly lemurs stay with their parents for up to two years following birth. (Harcourt and Thornback, 1990; Mittermeier, et al., 2006)

  • Parental Investment
  • altricial
  • female parental care
  • pre-fertilization
    • provisioning
    • protecting
      • female
  • pre-hatching/birth
    • provisioning
      • female
    • protecting
      • female
  • pre-weaning/fledging
    • provisioning
      • female
    • protecting
      • male
      • female
  • pre-independence
    • provisioning
      • male
      • female
    • protecting
      • male
      • female
  • extended period of juvenile learning

Lifespan/Longevity

Not much is known about the western woolly lemur's lifespan. Captured individuals have never survived for longer than a few days; the cause is hypothesized to be its highly selective folivorous diet. (Harcourt and Thornback, 1990)

Behavior

Western woolly lemurs live in family groups of 2 to 5 individuals; 3 is the most common number seen. Groups consist of a monogamous breeding pair with their young of up to two years. Family groups range around home territories of 1 to 2 hectares. Territory defense is rather lackluster, especially compared to closely related woolly lemurs in the eastern rain forests, Avahi laniger - western woolly lemurs tolerate much greater overlap between groups.

The western woolly lemurs, along with the other members of the genus Avahi, are the only nocturnal indriids. They are active at night and inactive during the day, although even at night, western woolly lemurs spend much of their time resting. Feeding is most common in the two hours after dusk and the two hours before dawn. Their seeming indolence is thought to be because their folivorous diet is not as energy-rich as other sources of food.

Like all indriids, western woolly lemurs are vertical clingers and leapers. Leaps have a mean length of 1.51 meters. Travel occurs most often during the first and last hour of the night. Family groups can cover large distances when traveling. (Harcourt and Thornback, 1990; Mittermeier, et al., 2006; Warren, 1997)

  • Range territory size
    10,000 to 20,000 m^2

Home Range

Not much is known about the home range of A. occidentalis. Territories are generally 1 to 2 hectares and are not strictly defended, in contrast to Avahi laniger. (Harcourt and Thornback, 1990; Mittermeier, et al., 2006)

Communication and Perception

Both sexes have scent glands on the neck that are used in olfactory communication. In addition to scent markings, the western woolly lemur uses several calls to communicate with its own group and with others. These include infant calls, which are plaintive, whistle-like noises used by the infant to attract its mother's attention. It also includes distant communication calls: a series of modulated, prolonged, high-pitched whistles. This call is used to help communicate territory borders and the receiver will reply with the same call. An alarm call varies with level of disturbance. In response to mild disturbance, it begins as a faint grunting sound, followed by a snorting sound. When highly disturbed, individuals use a loud, trembling call that sounds like "Ava Hy", hence the genus name. A cohesion call is a sudden, high-pitched call sounded when an individual is separated from another by 50 meters after an alarm. (Petter and Charles-Dominique, 1979)

Food Habits

Western woolly lemurs are highly selective folivores, only parts of the leaf blade are fed upon, never the midrib or the petiole. Targeted leaves are usually immature, although mature leaves will also be consumed. After studying leaves eaten by individuals, it was found that none contained a significant amount of alkaloids. At the same time, the species of plants consumed generally exhibit protein and sugar rich leaves. Individuals are very selective about their leaves; more than 20 species of plant have been reported as food sources, many of which are not commonly found in the forests of western Madagascar. Feeding, along with most other activities, occurs in the canopy, at heights ranging from 2 to 9 meters. Interestingly, when sharing habitat with Lepilemur species, which are also folivorous, A. occidentalis generally prevents Lepilemur from accessing better-quality food sources. (Ganzhorn, 1993; Ganzhorn, et al., 1985)

  • Plant Foods
  • leaves

Predation

The closely related A. laniger is a favored food of Malagasy raptors. It is reasonable to assume that the same raptors also prey on A. occidentalis. Their pelage acts as camouflage, blending into the trees it clings from. (Mittermeier, et al., 2006)

  • Anti-predator Adaptations
  • cryptic

Ecosystem Roles

Western woolly lemurs are most likely preyed upon by the raptors that co-occur with them. Where they co-occur with Lepilemur species, they restrict access to quality food resources for these Lepilemur species.

Little is known about lemur endoparasites, and, indeed, lemur parasites in general: the last great period of effort in naming and describing lemur parasites began around 1950 and petered out around the 1960's. Since then, studies have tended to focus more on the impact of parasites on their host species than the parasites themselves; A. occidentalis is no exception. Note: of the species listed, only Dipetalonema petteri has been found in A. occidentalis; the remainder were described in the closely related A. laniger and, due to the at-times questionable relationship between the two species, it is possible that they infect A. occidentalis as well.

An interesting note about the worm Dipetalonema petteri: although it was found in multiple necropsies of many lemur species, no males have ever been noted - only female worms have been recovered. The reasons behind this remain unknown. (Ganzhorn, 1993; Irwin and Raharison, 2009; Maa, 1969; Mittermeier, et al., 2006)

Commensal/Parasitic Species
  • Dipetalonema petteri
  • Allobosca crassipes
  • Pararhabdonema longistriata
  • Bertiella lemuriformis
  • Phtirpediculus avahidis

Economic Importance for Humans: Positive

Individually, western woolly lemurs have negligible positive impacts for humans, but the fascinating highly endemic Malagasy fauna, including other lemur species, are a major ecotourist attraction.

Economic Importance for Humans: Negative

Western woolly lemurs have no negative impact on humans.

Conservation Status

The IUCN lists A. occidentalis as endangered. Factors include its small range, fragmented populations, and recent declines in habitat and population. The most significant threat to the species is habitat destruction - mostly yearly burning of forest in order to generate new grazing land for cattle. It is also hunted to some degree. (Andrainarivo, et al., 2008; Mittermeier, et al., 2006)

Other Comments

Western woolly lemurs have variously been listed as the same species as the woolly lemur, Avahi laniger, considered a subspecies of A. laniger, or its own separate species. Most recently, it has been named a separate species, with the name Avahi occidentalis. (Harcourt and Thornback, 1990; Mittermeier, et al., 2006; Tattersall, 1982)

Contributors

Andrew Yu (author), Yale University, Eric Sargis (editor), Yale University, Rachel Racicot (editor), Yale University, Tanya Dewey (editor), University of Michigan-Ann Arbor.

Glossary

Ethiopian

living in sub-Saharan Africa (south of 30 degrees north) and Madagascar.

World Map

acoustic

uses sound to communicate

altricial

young are born in a relatively underdeveloped state; they are unable to feed or care for themselves or locomote independently for a period of time after birth/hatching. In birds, naked and helpless after hatching.

arboreal

Referring to an animal that lives in trees; tree-climbing.

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

chemical

uses smells or other chemicals to communicate

cryptic

having markings, coloration, shapes, or other features that cause an animal to be camouflaged in its natural environment; being difficult to see or otherwise detect.

ecotourism

humans benefit economically by promoting tourism that focuses on the appreciation of natural areas or animals. Ecotourism implies that there are existing programs that profit from the appreciation of natural areas or animals.

endothermic

animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.

female parental care

parental care is carried out by females

folivore

an animal that mainly eats leaves.

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

herbivore

An animal that eats mainly plants or parts of plants.

island endemic

animals that live only on an island or set of islands.

iteroparous

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

monogamous

Having one mate at a time.

motile

having the capacity to move from one place to another.

native range

the area in which the animal is naturally found, the region in which it is endemic.

nocturnal

active during the night

saltatorial

specialized for leaping or bounding locomotion; jumps or hops.

seasonal breeding

breeding is confined to a particular season

sedentary

remains in the same area

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

social

associates with others of its species; forms social groups.

tactile

uses touch to communicate

terrestrial

Living on the ground.

tropical

the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.

visual

uses sight to communicate

viviparous

reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.

References

2007. "Western Woolly Lemur" (On-line). The Primata. Accessed August 16, 2012 at http://www.theprimata.com/avahi_occidentalis.html.

Andrainarivo, C., V. Andriaholinirina, A. Feistner, T. Felix, J. Ganzhorn, N. Garbutt, C. Golden, B. Konstant, E. Louis Jr., D. Meyers, R. Mittermeier, A. Perieras, F. Princee, J. Rabarivola, B. Rakotosamimanana, H. Rasamimanana, J. Ratsimbazafy, G. Raveloarinoro, A. Razafimanantsoa, Y. Rumpler, C. Schwitzer, U. Thalmann, L. Wilmé, P. Wright. 2008. "The IUCN Red List of Threatened Species" (On-line). Avahi occidentalis. Accessed May 11, 2012 at http://www.iucnredlist.org/apps/redlist/details/2435/0.

Fleagle, J. 1999. Primate Adaptation and Evolution. USA: Academic Press.

Ganzhorn, J., J. Abraham, M. Razananhoera-Rakotomalal. 1985. Some aspects of the natural history and food selection of Avahi laniger. Primates, 26(4): 452-463.

Ganzhorn, J. 1993. Flexibility and Constraints of Lepilemur Ecology. Pp. 153-165 in P Kappeler, J Ganzhorn, eds. Lemur Social Systems and their Ecological Basis. USA: Plenum Press.

Garbutt, N. 2007. Mammals of Madagascar: A Complete Guide. New Haven, CT: Yale University Press.

Harcourt, C., J. Thornback. 1990. Lemurs of Madagascar and the Comoros: The IUCN Red Data Book. Gland, Switzerland: IUCN.

Irwin, M., J. Raharison. 2009. A Review of the Endoparasites of the Lemurs of Madagascar. Malagasy Nature, 2: 66-93. Accessed June 06, 2012 at http://www.sadabe.org/Irwin/articles/19_Irwin%20Raharison%202009.pdf.

Klopfer, P., K. Boskoff. 1979. Maternal Behavior in Prosimians. Pp. 123-156 in G Doyle, R Martin, eds. The Study of Prosimian Behavior. New York: Academic Press.

Maa, T. 1969. A Revised Checklist and Concise Host Index of Hippoboscidae (Diptera). Pacific Insects Monograph, 20: 261-299.

Mittermeier, R., W. Konstant, F. Hawkins, E. Louis, O. Langrand, J. Ratsimbazafy, R. Rasoloarison, J. Ganzhorn, S. Rajaobelina, I. Tattersall, D. Meyers. 2006. Lemurs of Madagascar. Colombia: Conservation International.

Petter, J., P. Charles-Dominique. 1979. Vocal Communication in Prosimians. Pp. 247-306 in G Doyle, R Martin, eds. The Study of Prosimian Behavior. New York: Academic Press.

Tattersall, I. 1982. The Primates of Madagascar. New York: Columbia University Press.

Thalmann, U. 2001. Food Resource Characteristics in Two Nocturnal Lemurs with Different Social Behavior: Avahi occidentalis and Lepilemur edwardsi. International Journal of Primatology, 22(2): 287-324.

Thalmann, U., T. Geissmann. 2000. Distribution and Geographic Variation in the Western Woolly Lemur (Avahi occidentalis) with Description of a New Species (A. unicolor). International Journal of Primatology, 21(6): 915-941.

Warren, R. 1997. Habitat Use and Support Preference of Two Free-Ranging Saltatory Lemurs (Lepilemur edwardsi and Avahi occidentalis). Journal of Zoology, 241(2): 325-341.