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Ring-tailed lemur

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Ring-tailed Lemur[1]
Scientific classification
Kingdom:
Animalia
Phylum:
Chordata
Class:
Mammalia
Order:
Primates
Family:
Lemuridae
Genus:
Lemur

Linnaeus, 1758
Species:
L. catta
Binomial name
Lemur catta
Linnaeus, 1758
Distribution of Lemur catta[3]

The Ring-tailed Lemur (Lemur catta) is a relatively large Strepsirhine primate, a lemur belonging to the family Lemuridae. The Ring-tailed Lemur is the only species within the monotypic genus Lemur and, like all other lemurs, is found only on the island of Madagascar. Known natively as Hira[3][4][5] (Malagasy) or Maki[3][5] (French, Malagasy), it inhabits gallery forests to spiny scrub in the southern regions of the island. In addition to being diurnal, it is also the most terrestrial of all lemurs.

The Ring-tailed Lemur is widely recognized and easily distinguished from other lemur species by its long, black and white ringed tail. It is highly social, as well as matriarchal, a trait uncommon in primates. Like other lemurs, this species relies strongly on its sense of smell, and marks its territory with scent glands. Furthermore, it is one of the most vocal primates, with numerous vocalizations, including group cohesion and alarm calls.[5]

Although threatened by habitat destruction and therefore listed as vulnerable by the IUCN Red List,[2] the Ring-tailed Lemur is the most populous lemur in zoos worldwide, with more than 1000 in captivity.[2][6] They reproduce readily in captivity and typically live up to 27 years (16 to 19 years in the wild).[7] Facilities actively involved in its conservation include the Duke Lemur Center in Durham, NC, the Myakka City Lemur Reserve (part of the Lemur Conservation Foundation) in Myakka City, FL, the Madagascar Fauna Group headquartered at the Saint Louis Zoo, and the Berenty Reserve in southern Madagascar.

Evolutionary history

No mammalian fossil record exists in Madagascar until recent times.[3] Consequently, little is known about the evolution of the Ring-tailed Lemur, let alone the order Lemuriformes, which comprises the entire endemic primate population of the island. However, chromosomal and molecular evidence strongly suggest that all lemurs are more closely related to each other than to other Strepsirrhine primates, which further suggests that a very small ancestral population came to Madagascar via a single rafting event between 47-57 million years ago.[3][8] Subsequent evolutionary radiation and speciation has created the diverse array of Malagasy lemurs seen today. As for the Ring-tailed Lemur, it is currently thought that it may share closer affinities to the bamboo lemurs of the genus Hapalemur.[3][8]

Taxonomic classification

The genus Lemur contains only one species, the Ring-tailed Lemur.

Changes in taxonomy

The Ring-tailed Lemur, along with brown lemurs and ruffed lemurs, were once grouped together in the genus Lemur. In fact, the Ring-tailed Lemur's skeleton is nearly indistinguishable from that of the brown lemurs.[8] However, in 1988, due to similarities between the Ring-tailed Lemur and the bamboo lemurs, particularly in regards to molecular evidence and similarities in the scent glands, the genus Lemur was split into the genera Lemur, Eulemur, and Varecia.[3][8] Consequently, the genus Lemur is now monotypic, containing only the Ring-tailed Lemur. Although not all authorities agree, the majority of the primatological community currently favors this classification.[3][9]

Anatomy and physiology

The Ring-tailed Lemur's tail is longer than its body

An adult Ring-tailed Lemur may reach a body length between 39 and 46 cm (15 and 18 in) and a weight between 2.3 and 3.5 kg (5.1 and 7.7 lb).[3] The Ring-tailed Lemur has a slender frame and narrow face, reminiscent of a vulpine muzzle. Like all lemurs, the Ring-tailed Lemur has hind limbs longer than its forelimbs. Females have two pairs of mammary glands, but only one pair is functional.[6]

Tail

The Ring-tailed Lemur's trademark, a long, bushy tail, is ringed in alternating black and white transverse stripes, numbering thirteen to fifteen each for both colors, and always ending in a black tip.[3][6][8] Its tail is longer than its body, measuring up to Template:Cm to in in length.[3] The tail is not prehensile and is only used for balance, communication, and group cohesion.[5]

Pelage and skin

The pelage is dense, with the ventral (chest) coat and throat being white or cream, and the dorsal (back) coat and neck being gray to rosy-brown. The crown is dark gray, while the ears and cheeks are white. The muzzle is dark grayish and the nose is black, and the eyes are encompassed by black lozenge-shaped patches.[3]

The skin is black in color, and visible on the nose, genitalia, and the palms and soles of the limbs. Like all lemurs, the Ring-tailed Lemur's fingers are slender, padded, and semi-dexterous with flat, human-like nails. Also in common with all lemurs, the Ring-tailed Lemur has a toilet-claw (sometimes referred to as a grooming claw) on the second toe of each hind limb specialized for grooming purposes.[10] Additionally, these primates groom orally by licking and tooth-scraping with narrow, procumbent lower incisors and canines, called a toothcomb.

Vision

The species' eyes can be a bright yellow or orange, which stay the same color from the time they are born. Unlike most diurnal primates, but like many prosimians, the Ring-tailed Lemur has a tapetum lucidum, or reflective layer on its eye, which enhances night vision.[6]

Scent glands

Furless scent glands are present on both males and females. Both genders have apocrine and sebaceous glands in their genital regions,[11] as well as antebrachial glands located on the inner surface of the forearm in proximity of the wrist. However, only the male has a horny spur that overlays this scent gland. The males also have brachial glands on the axillary surface of their shoulders.[6]

Ecology

The Ring-tailed Lemur is diurnal and semi-terrestrial. It is the most terrestrial of all lemur species, spending as much as 33% of its time on the ground. However, like other lemurs, it is also considerably arboreal, spending 23% of its time in the mid-level canopy, 25% in the upper-level canopy, 6% in the emergent layer, and 13% in small bushes. Troop travel is 70% terrestrial.[7]

Troop sizes range from 6 to 25, with 13 to 15 being the average and groups over 30 individuals being recorded.[8] Troop size, home range, and population density vary by region and food availability. Home range sizes range between 6 and 35 hectares (15 and 86 acres).[12] Troops of the Ring-tailed Lemur will maintain a territory, but overlap is often high. When encounters occur, they are agonistic. A troop will usually occupy the same part of its range for 3 or 4 days before moving. When it does move, the average traveling distance is 1 km (0.62 mi).[7] Population density ranges from 100 individuals per 1 km2 (0.39 sq mi) in dry forests to 250-600 individuals per km² in gallery and secondary forests.[3]

Geographic range and habitat

Found in southern and southwestern Madagascar and ranging further into highland areas than any other lemur, the Ring-tailed Lemur inhabits deciduous forests, dry scrub, montane humid forests, and gallery forests (forests along riverbanks). The Ring-tailed Lemur strongly favors gallery forests; such forests have now been cleared from much of Madagascar in order to create pasture for livestock.[3] Depending on location, temperatures within its geographic range can range between −7 °C (19 °F) to 48 °C (118 °F).[7]

This species is found as far east as Tôlanaro, north to Belo sur Tsiribihina, along the west coast, and inland towards the mountains of Andringitra on the southeastern plateau. It can be seen in the Berenty Reserve, Andohahela National Park, Isalo National Park, and Zombitse-Vohibasia National Park.[3]

Sympatric relations

The following lemur species can be found within the Ring-tailed Lemur geographic range:[7]

In Western Madagascar, sympatric Ring-tailed Lemurs and Red-fronted Brown Lemurs have been studied together. Little interaction takes place between the two species. Ring-tailed lemurs spend more time on the ground than the Brown Lemurs. Also, while the diets of the two species overlap, they eat the foods in different proportions, with the Ring-tailed Lemurs having a more varied diet.[13]

Behaviour

Ring-tailed Lemur eating the fruit (seed pod) of a tamarind tree

Diet

The Ring-tailed Lemur is an opportunistic omnivore, although it primarily eats fruits and leaves, particularly those of the tamarind tree (Tamarindus indica).[13] When available, tamarind can make up as much as 50% of the Ring-tailed Lemur's diet annually, especially during the dry, winter season. However, the Ring-tailed Lemur is known to eat from as many as 3 dozen different plant species and also includes flowers, herbs, bark, and sap in its diet. Additionally, it has been observed eating decayed wood, earth, spider webs, insect cocoons, arthropods (spiders, caterpillars, cicadas, and grasshoppers), and small vertebrates (birds and chameleons).[3] It becomes increasingly opportunistic during the dry season.

Social organization & interactions

The Ring-tailed Lemur lives in multi-male/multi-female troops, with a matriline as the core group. As with most lemurs, females socially dominate males in all circumstances, including feeding priority. Dominance is enforced by lunging, chasing, cuffing, grabbing, and biting. Young females do not inherit their mother's rank, while young males leave the troop between 3 and 5 years of age.[13] Both sexes have separate dominance hierarchies, with females having a distinct hierarchy and male rank being correlated with age. Generally, each troop has one to three central, high-ranking adult males, who interact with females more than other group males and lead the troop procession with high-ranking females. Lower ranking males are typically recently transferred males, old males, or young adult males that have not yet left their natal group. They tend to be marginalized from group activity, staying at the periphery of the group.[7]

For males, social structure changes can be seasonal, with males immigrating between groups during the 6-month period between December through May. Established males transfer every 3.5 years,[13] although young males may transfer every 1.4 years. Group fission occurs when groups get too large and resources become scarce.[7]

Ring-tailed Lemurs sunning

In the mornings, the Ring-tailed Lemur sunbathes to warm itself. It faces the sun, sitting in what is frequently described as a "sun-worshipping" posture or Lotus position. However, it sits with its legs extended outward, not cross-legged, and will often support itself on nearby branches. Sunning is often a group activity, particularly during the cold mornings. At night, troops will split into sleeping parties, huddling closely together to keep warm. Groups of huddled Ring-tailed Lemurs are popularly referred to as lemur balls.

Despite being primarily quadrupedal, the Ring-tailed Lemur can rear up and balance on its hind legs, usually for aggressive displays. When threatened, the Ring-tailed Lemur has been known to strike out with its short nails in a behaviour termed jump fighting. This action is extremely rare outside of the breeding season when tensions are high and competition for access to mates is intense. Other aggressive behaviours include a threat-stare, used to intimidate or start a fight, and a submissive gesture known as pulled-back lips.[7]

Border disputes with rival troops occur occasionally, and it is the dominant female's responsibility to defend the troop's home range. Agonistic encounters include staring, lunging approaches, and occasional physical aggression, and conclude with troop members retreating toward the center of the home range.[7]

Ring-tailed Lemur scent-marking using anogenital scent glands

Olfactory communication

Like most other prosimians, olfactory communication is critically important for the Ring-tailed Lemur. Males and females scent mark both vertical and hortizontal surfaces at the overlaps in their home ranges using their anogenital scent glands. In order to mark vertical surfaces, the Ring-tailed Lemur will perform a handstand, grasping the highest point possible with its feet while it applies its scent.[14] Use of scent marking varies by age, sex and social status.[15] Male lemurs use their antebrachial and brachial glands to demarcate territories and maintain intragroup dominance hierarchies. The thorny spur that overlays the antebrachial gland on each wrist is scraped against tree trunks to create grooves anointed with their scent. This is known as spur-marking.[14]

In displays of aggression, males will engage in a social display behaviour called stink fighting, which involves impregnating their tails with secretions from the antebrachial and brachial glands, and then waving the scented tail at male rivals.[16] Males will also occasionally wave their scented tails at females as a form of sexual overture; this usually results in the female cuffing or biting the male, and elicits subordinate vocalizations from the would-be paramour.

Auditory communication

Main article: Calls of the Ring-tailed Lemur
This article includes inline links to audio files. If you have trouble playing the files, see Wikipedia Media help.

The Ring-tailed Lemur has a complex array of distinct vocalizations used to maintain group cohesion during foraging and alert group members to the presence of a predator. Calls range from simple, such as the purr Lemur_catta--purr1.oggⓘ, which expresses contentment, to complex, such as the sequence of clicks, close-mouth click series (CMCS), open-mouth click series (OMCS), and yaps Lemur_catta--click_series_&_yaps.oggⓘ, which is used during predator mobbing.[17] Some calls have variants and undergo transitions between variants and/or other calls, such as an infant "whit" (distress call) transitioning from variant 1 to variant 2 Lemur_catta--infant_whits_transition_var1-var2.oggⓘ.[17]

The most commonly heard vocalizations of the Ring-tailed Lemur are the moan Lemur_catta--moan1.oggⓘ (low-to-moderate arousal, group cohesion), early-high wail Lemur_catta--early-high_wails1.oggⓘ (moderate-to-high arousal, group cohesion), and clicks Lemur_catta--clicks1.oggⓘ ("location marker" to draw attention).[17]


A female carrying new-born twins

Breeding and reproduction

The Ring-tailed Lemur is polygynous, although one central male typically breeds with more females than the others do. Fighting is most common during the breeding season.[18] When receptive, females will present their backside, lift their tail, and look at her desired male over her shoulder. Males will typically inspect the female's genitals to determine receptiveness. Although females typically mate within their troop, they will occasionally seek out males from outside their troop.[13]

The breeding season runs from mid-April to mid-May, with estrus lasting approximately 4-6 hours.[8] Females mate with multiple males during their estrus period.[13] Gestation lasts for about 135 days, with birth of the young generally occurring in September, but occasionally in October.[8] In the wild, one offspring is the norm, although twins may occur.[8] Ring-tailed Lemur infants have a birth weight of 70 g (2.5 oz) and are carried ventrally (on the chest) for the first 1–2 weeks, then dorsally (on the back).[8] The young lemurs begin to eat solid food after two months and are fully weaned after five months. Sexual maturity is reached between 2.5–3 years.[18] Male involvement in infant rearing is limited, although the entire troop, regardless of age or sex, can be seen caring for the young. Alloparenting between troop females can be seen, with instances of kidnapping having been reported. Infanticide by males also occurs occasionally.[7] Due to harsh environmental conditions, accidents (such as falls), and predation, infant mortality can be as high as 50% within the first year, with as few as 30% reaching adulthood.[8]

Threats in the wild

Human activity is the greatest threat to Ring-tailed Lemurs. Habitat destruction is the greatest concern, with much of their range being cleared through annual burning to create pasture for livestock. Furthermore, overgrazing and the felling of trees for charcoal production are having a severe impact. This species is also hunted for food (bush meat) and kept as pets.[19]

The Ring-tailed Lemur has both natural and introduced predators. Native predators include the Fossa (Cryptoprocta ferox), the Madagascar Harrier-hawk (Polyboroides radiatus), the Madagascar Buzzard (Buteo brachypterus), and the Madagascar Ground Boa (Boa madagascariensis). Introduced predators include the Small Indian Civet (Viverricula indica), the Domestic Cat, and the Domestic Dog.[3]

The Ring-tailed Lemur has been popularized in the Animal Planet television series Lemur Kingdom and in the animated film Madagascar (2005). The television series depicts real events in the lives of wild Ring-tailed Lemurs, whereas the animated film depicts anthropomorphic representations, with lemurs talking, singing, and dancing.

See also

References

  1. ^ Groves, C. P. (2005). Wilson, D. E.; Reeder, D. M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Baltimore: Johns Hopkins University Press. p. 117. ISBN 0-801-88221-4. OCLC 62265494.
  2. ^ a b c Template:IUCN2006 Listed as Vulnerable (VU A1c v2.3)
  3. ^ a b c d e f g h i j k l m n o p q Mittermeier, R.A. (2006). Lemurs of Madagascar (2nd Edition ed.). Conservation International. pp. 23–26, 237–249. ISBN 1-881173-88-7. {{cite book}}: |edition= has extra text (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)
  4. ^ McGinley, M (2008-08-21). "Primate Factsheets: Madagascar spiny thickets". Encyclopedia of Earth. Retrieved 2008-08-30. {{cite web}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  5. ^ a b c d "Ringtailed Lemur". Duke Lemur Center. Retrieved 2008-08-30.
  6. ^ a b c d e "Ring-tailed Lemur, Lemur catta fact sheet". San Diego Zoo Library. 2003. Retrieved 2008-08-30.
  7. ^ a b c d e f g h i j Cawthon Lang, KA (2005-07-21). "Primate Factsheets: Ring-tailed lemur (Lemur catta) Taxonomy, Morphology, & Ecology". Wisconsin Primate Research Center (WPRC). Retrieved 2008-04-18.
  8. ^ a b c d e f g h i j k Garbutt, N. (2007). Mammals of Madagascar, A Complete Guide. A&C Black Publishers. pp. 85–86, 146–148. ISBN 978-0-300-12550-4.
  9. ^ Groves, C. P. (2005). Wilson, D. E.; Reeder, D. M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Baltimore: Johns Hopkins University Press. p. Lemur. ISBN 0-801-88221-4. OCLC 62265494.
  10. ^ "Lemur Basics". Retrieved 2008-08-28.
  11. ^ Scordato, E.S., Dubay, G., & Drea, C.M. (2007). "Chemical Composition of Scent Marks in the Ringtailed Lemur (Lemur catta): Glandular Differences, Seasonal Variation, and Individual Signatures". Chemical Senses. 32: 493–504. doi:10.1093/chemse/bjm018.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  12. ^ Gould, L. and Sauther, M. (2007). "Lemuriformes". In Campbell, C., Fuentes, A., MacKinnon, K., Panger, M., & Bearder, S. (ed.). Primates in Perspective. p. 53. ISBN 978-0-19-517133-4.{{cite book}}: CS1 maint: multiple names: authors list (link)
  13. ^ a b c d e f Sussman, R. (1999). Primate Ecology and Social Structure Volume 1: Lorises, Lemurs and Tarsiers. p. 154-173. ISBN 0-536-02256-9.
  14. ^ a b Cawthon Lang, KA (2005-07-21). "Primate Factsheets: Ring-tailed lemur (Lemur catta) Behavior". Wisconsin Primate Research Center (WPRC). Retrieved 2008-08-28.
  15. ^ Gouzoules, H. and Gouzoules, S. (2007). "The Conundrum of Communication". In Campbell, C., Fuentes, A., MacKinnon, K., Panger, M., & Bearder, S. (ed.). Primates in Perspective. p. 624. ISBN 978-0-19-517133-4.{{cite book}}: CS1 maint: multiple names: authors list (link)
  16. ^ Rowe, N. (1996). The Pictorial Guide to the Living Primates. p. 38. ISBN 0-9648825-0-7.
  17. ^ a b c Macedonia, Joseph M. (1993). "The Vocal Repertoire of the Ringtailed Lemur (Lemur catta)". Folia Primatologica. 61: 186–217.
  18. ^ a b Anderson, R (1999). "Lemur catta". Animal Diversity Web. Retrieved 2008-04-18.
  19. ^ Cawthon Lang, KA (2005-07-21). "Primate Factsheets: Ring-tailed lemur (Lemur catta) Conservation". Wisconsin Primate Research Center (WPRC). Retrieved 2008-08-28.
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