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<title>Ring-tailed Lemur</title>
<authors>Wikipedia</authors>
<publisher>wikipedia.org</publisher>
<publishedplace>network</publishedplace>
<publishedyear>2010</publishedyear>
<edition></edition>
<category>IUCN Red List near threatened species</category>
<section id="23105" fatherid="0" title="" lv="" priority="-1" level="1">
The Ring-tailed Lemur (Lemur catta) is a large Strepsirhine primate and the most recognized lemur due to its long, black and white ringed tail. It belongs to Lemuridae, one of four lemur families. It is the only member of the Lemur genus. Like all lemurs it is endemic to the island of Madagascar. Known locally as Hira (Malagasy), Maky (Malagasy), or Maki (French),Mittermeier, pp. 246 it inhabits gallery forests to spiny scrub in the southern regions of the island. It is omnivorous and the most terrestrial of lemurs. The animal is diurnal, being active exclusively in daylight hours.
The Ring-tailed Lemur is highly social, living in groups of up to 30 individuals.  It is also female dominant, a trait common among lemurs. To keep warm and reaffirm social bonds, groups will huddle together forming a lemur ball. The Ring-tailed Lemur will also sunbathe, sitting up facing its underside, with its thinner white fur towards the sun. Like other lemurs, this species relies strongly on its sense of smell and marks its territory with scent glands.  The males perform a unique scent marking behavior called spur marking and will participate in stink fights by impregnating their tail with their scent and wafting it at opponents.
As one of the most vocal primates, the Ring-tailed Lemur utilizes numerous vocalizations including group cohesion and alarm calls.  Despite the lack of a large brain (relative to Simiiform primates) experiments have shown that the Ring-tailed Lemur can organize sequences, understand basic arithmetic operations and preferentially select tools based on functional qualities.
Despite being listed as Near Threatened by the IUCN Red List and suffering from habitat destruction, the Ring-tailed Lemur reproduces readily in captivity and is the most populous lemur in zoos worldwide, numbering more than 2000 individuals. It typically lives 16 to 19 years in the wild and 27 years in captivity.

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<section id="23106" fatherid="0" title="Etymology" lv="" priority="-2" level="1">
The term lemur was selected by early biologists because the calls of some elusive lemur species brought to mind the cries of the spirits of the dead, or lemures, from  Roman mythology.Garbutt, pp. 85-86  The species name, catta, comes from the similarity between the Ring-tailed Lemur's purring vocalization and that of the Domestic Cat.
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<section id="23107" fatherid="0" title="Evolutionary history" lv="" priority="3" level="1">
<section id="23108" fatherid="23107" title="" lv="" priority="-1" level="2">
All mammalian fossils from Madagascar come from recent times.Mittermeier, pp. 23-26  Thus, little is known about the evolution of the Ring-tailed Lemur, let alone the infraorder Lemuriformes, which comprises the entire endemic primate population of the island.  However, chromosomal and molecular evidence suggest that lemurs are more closely related to each other than to other Strepsirrhine primates.  For this to have happened, it is thought that a very small ancestral population came to Madagascar via a single rafting event between 50 and 80 million years ago. Subsequent evolutionary radiation and speciation has created the diversity of Malagasy lemurs seen today.  The Ring-tailed Lemur itself is thought to share closer affinities than the rest of its subfamily, Lemurinae, to the bamboo lemurs of the genus Hapalemur,Garbutt, pp. 146-148Mittermeier, pp. 237 which may be a sister group of the family Lemuridae.
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<section id="23109" fatherid="23107" title="Taxonomic classification" lv="" priority="2" level="2">
<section id="23110" fatherid="23109" title="" lv="" priority="-1" level="3">
The genus Lemur contains only one species, the Ring-tailed Lemur.
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<section id="23111" fatherid="23109" title="Changes in taxonomy" lv="" priority="-2" level="3">
The Ring-tailed Lemur shares many similarities with the members of its subfamily, Lemurinae, and its skeleton is nearly indistinguishable from that of the brown lemurs.  For these reasons, the Ring-tailed Lemur, brown lemurs and ruffed lemurs were once grouped together in the genus Lemur.  However, ruffed lemurs were reassigned to the genus Varecia in 1962, and due to similarities between the Ring-tailed Lemur and the bamboo lemurs, particularly in regards to molecular evidence and scent glands similarities, the brown lemurs were moved to the genus Eulemur in 1988. 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.
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<section id="23112" fatherid="0" title="Anatomy and physiology" lv="" priority="-4" level="1">
An adult Ring-tailed Lemur may reach a body length between  and a weight between .Mittermeier, pp. 246-249  The species has a slender frame and narrow face, reminiscent of a vulpine muzzle. Like all lemurs, its hind limbs are longer than its forelimbs. Females have two pairs of mammary glands, but only one pair is functional.
Furless scent glands are present on both males and females. Both genders have apocrine and sebaceous glands in their genital regions, 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.
The Ring-tailed Lemur's trademark, a long, bushy tail, is ringed in alternating black and white transverse stripes, numbering 13 to 15 each for both colors, and always ending in a black tip. Its tail is longer than its body, measuring up to  in length. The tail is not prehensile and is only used for balance, communication, and group cohesion.
The pelage, or fur, is dense. The ventral (chest) coat and throat are white or cream, and the dorsal (back) coat is 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.
The black skin is visible on the nose, genitalia, and the palms and soles of the limbs. The Ring-tailed Lemur shares several adaptations with other lemurs.  Its fingers are slender, padded, and semi-dexterous with flat, human-like nails. It grooms orally by licking and tooth-scraping with narrow, procumbent lower incisors and canines, called a toothcomb. Lastly, it has a toilet-claw (sometimes referred to as a grooming claw) on the second toe of each hind limb specialized for personal grooming, specifically to rake through fur that is unreachable by the mouth and toothcomb.
The species' eyes can be a b yellow or orange. Unlike most diurnal primates, but like all strepsirhine primates, the Ring-tailed Lemur has a tapetum lucidum, or reflective layer behind the retina of the eye, that enhances night vision.

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<section id="23113" fatherid="0" title="Ecology" lv="" priority="-5" level="1">
<section id="23114" fatherid="23113" title="" lv="" priority="-1" level="2">
The Ring-tailed Lemur is diurnal and semi-terrestrial. It is the most terrestrial of lemur species, spending as much as 33% of its time on the ground. However it is still 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.
Troop size, home range, and population density vary by region and food availability. Troops typically range in size from 6 to 25, although troops with over 30 individuals have been recorded. The average troop contains 13 to 15 individuals. Home range sizes varies between . Troops of the Ring-tailed Lemur will maintain a territory, but overlap is often high. When encounters occur, they are agonistic, or hostile in nature. A troop will usually occupy the same part of its range for three or four days before moving. When it does move, the average traveling distance is . Population density ranges from 100 individuals per  in dry forests to 250–600 individuals per km2 in gallery and secondary forests.
The Ring-tailed Lemur has both native 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.

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<section id="23115" fatherid="23113" title="Geographic range and habitat" lv="" priority="2" level="2">
Endemic to southern and southwestern Madagascar, the Ring-tailed Lemur ranges further into highland areas than other lemurs.  It inhabits deciduous forests, dry scrub, montane humid forests, and gallery forests (forests along riverbanks). It strongly favors gallery forests, but such forests have now been cleared from much of Madagascar in order to create pasture for livestock. Depending on location, temperatures within its geographic range can vary between  and .
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.

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<section id="23116" fatherid="23113" title="Sympatric relations" lv="" priority="3" level="2">
The following lemur species are sympatric with the Ring-tailed Lemur, meaning they are related species found within its geographic range:
In Western Madagascar, sympatric Ring-tailed Lemurs and Red-fronted Brown Lemurs have been studied together. Little interaction takes place between the two species. The Ring-tailed Lemur spends more time on the ground than the Red-fronted Brown Lemur. While the diets of the two species overlap, they eat in different proportions since the Ring-tailed Lemur has a more varied diet.

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<section id="23117" fatherid="0" title="Behavior" lv="" priority="6" level="1">
<section id="23118" fatherid="23117" title="Diet" lv="" priority="1" level="2">
The Ring-tailed Lemur is an opportunistic omnivore primarily eating fruits and leaves, particularly those of the tamarind tree (Tamarindus indica), known natively as kily. When available, tamarind makes up as much as 50% of the diet, especially during the dry, winter season. The Ring-tailed Lemur eats from as many as three dozen different plant species, and its diet includes flowers, herbs, bark and sap. It has been observed eating decayed wood, earth, spider webs, insect cocoons, arthropods (spiders, caterpillars, cicadas and grasshoppers) and small vertebrates (birds and chameleons). During the dry season it becomes increasingly opportunistic.
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<section id="23119" fatherid="23117" title="Social systems" lv="" priority="2" level="2">
<section id="23120" fatherid="23119" title="" lv="" priority="-1" level="3">
Troops are classified as multi-male/multi-female, 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 and young males leave the troop between three and five years of age.  Both sexes have separate dominance hierarchies; females have a distinct hierarchy while male rank is correlated with age. 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. Recently transferred males, old males or young adult males that have not yet  their natal group are often lower ranking. Staying at the periphery of the group they tend to be marginalized from group activity.
For males, social structure changes can be seasonal. During the six month period between December and May a few males immigrate between groups. Established males transfer every 3.5 years, although young males may transfer every 1.4 years. Group fission occurs when groups get too large and resources become scarce.
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. A group of huddled Ring-tailed Lemurs is referred to as a lemur ball.
Despite being quadrupedal the Ring-tailed Lemur can rear up and balance on its hind legs, usually for aggressive displays. When threatened the Ring-tailed Lemur may jump in the air and strike out with its short nails and sharp upper canine teeth in a behaviour termed jump fighting. This is extremely rare outside of the breeding season when tensions are high and competition for 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.
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.

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<section id="23121" fatherid="23119" title="Olfactory communication" lv="" priority="2" level="3">
Olfactory communication is critically important for prosimians like the Ring-tailed Lemur. Males and females scent mark both vertical and horizontal surfaces at the overlaps in their home ranges using their anogenital scent glands. The Ring-tailed Lemur will perform a handstand to mark vertical surfaces, grasping the highest point with its feet while it applies its scent. Use of scent marking varies by age, sex and social status. 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.
In displays of aggression, males engage in a social display behaviour called stink fighting, which involves impregnating their tails with secretions from the antebrachial and brachial glands and waving the scented tail at male rivals. During the mating season, males 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.

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<section id="23122" fatherid="23119" title="Auditory communication" lv="" priority="-3" level="3">
The Ring-tailed Lemur is one of the most vocal primates and 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 to complex.  An example of a simple call is the purr (), which expresses contentment.  A complex call is the sequence of clicks, close-mouth click series (CMCS), open-mouth click series (OMCS) and yaps () used during predator mobbing.  Some calls have variants and undergo transitions between variants, such as an infant "whit" (distress call) transitioning from one variant to another ().
The most commonly heard vocalizations are the moan () (low-to-moderate arousal, group cohesion), early-high wail () (moderate-to-high arousal, group cohesion), and clicks () ("location marker" to draw attention).

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<section id="23123" fatherid="23117" title="Breeding and reproduction" lv="" priority="3" level="2">
The Ring-tailed Lemur is polygynous, although the dominant male in the troop typically breeds with more females than other males. Fighting is most common during the breeding season. A receptive female may initiate mating by presenting her backside, lifting her tail and looking at the desired male over her shoulder. Males may inspect the female's genitals to determine receptiveness. Females typically mate within their troop, but may seek outside males.
The breeding season runs from mid-April to mid-May. Estrus lasts 4 to 6 hours, and females mate with multiple males during this period.  Within a troop, females stagger their receptivity so that each female comes into season on a different day during the breeding season, reducing competition for male attention. Gestation lasts for about 135 days, and parturition occurs in September or occasionally October. In the wild one offspring is the norm, although twins may occur. Ring-tailed Lemur infants have a birth weight of  and are carried ventrally (on the chest) for the first 1 to 2 weeks, then dorsally (on the back).
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 and 3 years. 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 has been reported. Kidnapping by females and infanticide by males also occur occasionally. Due to harsh environmental conditions, predation and accidents such as falls infant mortality can be as high as 50% within the first year and as few as 30% may reach adulthood.  The Ring-tailed Lemur can go on to live 16 to 19 years (27 years in captivity).

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<section id="23124" fatherid="23117" title="Cognitive abilities and tool use" lv="" priority="4" level="2">
Historically, the studies of learning and cognition in non-human primates have focused on simians (monkeys and apes), while strepsirrhine primates, such as the Ring-tailed Lemur and its allies, have been overlooked and popularly dismissed as unintelligent. A couple of factors stemming from early experiments have played a role in the development of this assumption. First, the experimental design of older tests may have favored the natural behavior and ecology of simians over that of strepsirrhines, making the experimental tasks inappropriate for lemurs. For example, simians are known for their manipulative play with non-food objects, whereas lemurs are only known to manipulate non-food objects in captivity.  This behaviour is usually connected with food association. Also, lemurs are known to displace objects with their nose or mouth more so than with their hands.  Therefore, an experiment requiring a lemur to manipulate an object without prior training would favor simians over strepsirrhines. Second, individual Ring-tailed Lemurs accustomed to living in a troop may not respond well to isolation for laboratory testing. Past studies have reported hysterical behaviour in such scenarios. As a result of these early studies lemurs were often omitted from further research.
The notion that lemurs are unintelligent has been perpetuated by the view that the neocortex ratio (as a measure of brain size) indicates intelligence. In fact, primatologist Alison Jolly noted early in her academic career that some lemur species, such as the Ring-tailed Lemur, have evolved a social complexity similar to that of cercopithecine monkeys, but not the corresponding intelligence. After years of observations of wild Ring-tailed Lemur populations at the Berenty Reserve in Madagascar and as well as baboons in Africa, she more recently concluded that this highly social lemur species does not demonstrate the equivalent social complexity of cercopithecine monkeys, despite general appearances.
Regardless, research has continued to illuminate the complexity of the lemur mind, with emphasis on the cognitive abilities of the Ring-tailed Lemur. As early as the mid-1970s, studies had demonstrated that they could be trained through operant conditioning using standard schedules of reinforcement. The species has been shown to be capable of learning pattern, bness and object discrimination, skills common among vertebrates. The Ring-tailed Lemur has also been shown to learn a variety of complex tasks often equaling, if not exceeding, the performance of simians.
More recently, research at the Duke Lemur Center has shown that the Ring-tailed Lemur can organize sequences in memory and retrieve ordered sequences without language. The experimental design demonstrated that the lemurs were using internal representation of the sequence to guide their responses and not simply following a trained sequence, where one item in the sequence cues the selection of the next. But this is not the limit of the Ring-tailed Lemur's reasoning skills. Another study, performed at the Myakka City Lemur Reserve, suggests that this species along with several other closely related lemur species understand simple arithmetic operations.
Since tool use is considered to be a key feature of primate intelligence, the apparent lack of this behavior in wild lemurs, as well as the lack of non-food object play, has helped reinforce the perception that lemurs are less intelligent than their simian cousins. However, another study at the Myakka City Lemur Reserve examined the representation of tool functionality in both the Ring-tailed Lemur and the Common Brown Lemur and discovered that, like monkeys, they utilized tools with functional properties (e.g., tool orientation or ease of use) instead of tools with nonfunctional features (e.g., color or texture). Although the Ring-tailed Lemur may not use tools in the wild, it can not only be trained to use a tool, but will preferentially select tools based on their functional qualities. Therefore, the conceptual competence to utilize a tool may have been present in the common primate ancestor, even though the use of tools may not have appeared until much later.

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<section id="23125" fatherid="0" title="Conservation status" lv="" priority="7" level="1">
In addition to being listed as Near Threatened in 2008 by the IUCN, the Ring-tailed Lemur has been listed since 1977 by CITES under Appendix I, which makes trade of wild-caught specimens illegal. Although there are more endangered species of lemur, the Ring-tailed Lemur is considered a flagship species due to its recognizability.
Three factors threaten Ring-tailed Lemurs. First and foremost is habitat destruction.  Starting nearly 2,000 years ago with the introduction of humans to the island, forests have been cleared to produce pasture and agricultural land.  Extraction of hardwoods for fuel and lumber, as well mining and overgrazing, have also taken their toll.  Today, it is estimated that 90% of Madagascar's original forest cover has been lost.Mittermeier, pp. 57  Rising populations have created even greater demand in the southwest portion of the island for fuel wood, charcoal, and lumber.  Fires from the clearing of grasslands, as well as slash-and-burn agriculture destroy forests.  Another threat to the species is harvesting either for food (bush meat) or pets.  Finally, periodic drought common to southern Madagascar can impact populations already in decline.  In 1991 and 1992, for example, a severe drought caused an abnormally high morality rate among infants and females at the Beza Mahafaly Special Reserve.  Two years later, the population had declined by 31% and took nearly four years to start to recover.
The Ring-tailed Lemur resides in several protected areas within its range, each offering varying levels of protection.  At the Beza Mahafaly Special Reserve, a holistic approach to in situ conservation has been taken.  Not only does field research and resource management involve international students and local people (including school children), livestock management is used at the peripheral zones of the reserve and ecotourism benefits the local people.
Outside of its diminishing habitat and other threats, the Ring-tailed Lemur reproduces readily and has fared well in captivity.  For this reason, along with its popularity, it has become the most populous lemur in zoos worldwide, with more than 2000 in captivity.  Ex situ facilities actively involved in the conservation of the Ring-tailed Lemur include the Duke Lemur Center in Durham, NC, the Lemur Conservation Foundation in Myakka City, FL and the Madagascar Fauna Group headquartered at the Saint Louis Zoo.  Due to the high success of captive breeding, reintroduction is a possibility if wild populations were to crash.  Although experimental releases have met success on St. Catherines Island in Georgia, demonstrating that captive lemurs can readily adapt to their environment and exhibit a full range of natural behaviors, captive release is not currently being considered.
Ring-tailed Lemur populations can also benefit from drought intervention, due to the availability of watering troughs and introduced fruit trees, as seen at the Berenty Private Reserve in southern Madagascar.  However, these interventions are not always seen favorably, since natural population fluctuations are not permitted.  The species is thought to have evolved its high fecundity due to its harsh environment; therefore, interfering with this natural cycle could significantly impact the gene pool.

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<section id="23126" fatherid="0" title="Cultural references" lv="" priority="-8" level="1">
The Ring-tailed Lemur, known locally as Hira (Malagasy), Maky (Malagasy), or Maki (French), has been popularized in Western culture by the Animal Planet television series Lemur Kingdom (United States) and Lemur Street (United Kingdom), as well as by the character King Julien in the animated Madagascar film series and spin-off TV series. Lemur Street depicts real events in the lives of wild Ring-tailed Lemurs, whereas the Madagascar films and spin-off series depict anthropomorphic representations, such as lemurs talking, singing, and dancing.  The Ring-tailed Lemur was also the focus of the 1996 Nature documentary A Lemur's Tale, which was filmed at the Berenty Reserve and followed a troop of lemurs.  The troop included a unique infant named Sapphire, who was nearly albino, with white fur, sparkling blue eyes, and the characteristic ringed tail.
This species also played a role in the 1997 comedy film Fierce Creatures, starring John Cleese, who has a passion for lemurs.  In fact, John Cleese also hosted the 1998 BBC documentary In the Wild: Operation Lemur with John Cleese.

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<section id="23127" fatherid="0" title="Notes" lv="" priority="-9" level="1">
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<section id="23128" fatherid="0" title="References" lv="" priority="-10" level="1">
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<section id="23129" fatherid="0" title="External links" lv="" priority="-11" level="1">
Category:Lemurs


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