The ring-tailed lemur (Lemur catta) is a medium- to large-sized primate that belongs to the strepsirrhine group, which includes animals with wet noses. It is the most well-known lemur species because of its long, black-and-white, ringed tail. This animal is part of the Lemuridae family, one of five lemur families, and is the only member of the Lemur genus. Like all lemurs, it lives only on the island of Madagascar, where it is classified as endangered. Locally, it is called maky or hira in the Malagasy language. It lives in different types of habitats, including forests near rivers and dry, spiny areas in the south of the island. It eats both plants and animals, and it is the lemur species best suited to living on the ground.
Ring-tailed lemurs live in groups called "troops" that can include up to 30 individuals. They are a female-dominant species, which is common among lemurs. To stay warm and strengthen group bonds, they huddle together. Grooming each other is an important part of their social behavior, helping to remove insects and reinforce relationships. These lemurs are active only during the day, a trait called diurnal. Because of this, they often sit on their tails in the sun, exposing their white belly fur. They may also hold their palms up and close their eyes while sunbathing. Like other lemurs, they rely heavily on their sense of smell. They mark their territory using scent glands near their eyes and near their anus. Males perform a special marking behavior called spur-marking and may "waft" their tails to spread pheromones during confrontations. Both males and females mark trees, logs, rocks, and other objects by rubbing their faces and bodies on them, similar to how cats mark their territory.
The ring-tailed lemur is one of the most vocal primates. It uses many sounds to communicate, such as calling to keep the group together or warning of dangers. Studies have shown that despite having a smaller brain compared to other primates, this species can arrange items in order, understand simple math, and choose tools based on how useful they are.
Although ring-tailed lemurs adapt well to living in captivity and are the most commonly seen lemur in zoos worldwide (with more than 2,000 raised in captivity), their wild population is endangered due to habitat loss, hunting for bushmeat, and the pet trade. As of early 2017, their wild population was estimated to be as low as 2,000 individuals. Local farmers and logging industries in Madagascar often use slash-and-burn techniques to clear land for farming and livestock, which contributes to the loss of their natural habitat.
Etymology
The word "lemur" was first used to describe slender lorises, but later it became the name for primates found only in Madagascar. These animals are now called lemurs. The name comes from the Latin word "lemures," which means ghosts or spirits that were driven away during a festival in ancient Rome. Carl Linnaeus, the scientist who named the slender loris, chose the name because the animal is active at night and moves slowly. He was inspired by writings from the Roman poets Virgil and Ovid, who described similar ideas. Some people incorrectly think Linnaeus named the animal because of its ghost-like appearance, glowing eyes, or eerie sounds. However, this is not true because the name was first used for slender lorises from India, not for lemurs. The species name "catta" refers to the ring-tailed lemur's cat-like look. Its sound, which is similar to a cat's purr, also helps explain this name.
After Linnaeus described the species, the name "ring-tailed maucauco" was first written in 1771 by Thomas Pennant, a Welsh naturalist. He noted the animal's long, banded tail. At that time, "maucauco" was a common name for lemurs. The widely used English name "ring-tailed lemur" was first used by George Shaw in a scientific book published between 1792 and 1796. This book illustrated specimens from the Leverian collection.
Evolutionary history
All mammal fossils found on Madagascar are from recent times. This means scientists know little about how the ring-tailed lemur evolved, or about the rest of the lemur group, which includes all primates that live only on the island. However, studies of chromosomes and molecules suggest that lemurs are more closely related to each other than to other types of primates called strepsirrhines. For this to happen, scientists think a small group of ancestors reached Madagascar through one event, such as floating on a piece of land, between 50 and 80 million years ago. Over time, these ancestors diversified and gave rise to the many types of lemurs found today.
Analysis of amino acid sequences shows that the family Lemuridae split into different groups around 26.1 million years ago, with a possible range of 3.3 million years. Studies of mitochondrial DNA suggest a slightly earlier split, around 24.9 million years ago, with a possible range of 3.6 million years. The ruffed lemurs were the first group to separate from the rest of the family, a finding supported by DNA and chromosome studies. Molecular data also show that true lemurs (Eulemur) are closely related to two other lemur groups, Lemur and Hapalemur, but not as closely to each other.
The ring-tailed lemur is thought to be more closely related to bamboo lemurs (Hapalemur) than to other lemur groups. This idea is supported by comparisons of communication, chromosomes, genetics, and physical traits like scent glands. However, other studies of immunity and some physical traits do not support this close relationship. For example, bamboo lemurs have short snouts, while the ring-tailed lemur and most other lemurs have long snouts. Differences in the structure of the eye sockets and face suggest that the ring-tailed lemur and true lemurs developed long snouts independently.
The relationship between the ring-tailed lemur and bamboo lemurs is unclear. Some genetic studies suggest that bamboo lemurs split from the ring-tailed lemur, keeping the group as a single evolutionary line (monophyletic) and supporting the current classification into two genera. Other studies suggest the ring-tailed lemur might be part of the bamboo lemur group, which would require splitting the bamboo lemur species into a new genus called Prolemur.
The ring-tailed lemur has 56 chromosomes. Four are metacentric (arms of nearly equal length), four are submetacentric (arms of unequal length), and 46 are acrocentric (the short arm is barely visible). The X chromosome is metacentric, and the Y chromosome is acrocentric.
In 1754, Linnaeus first used the name Lemur to describe a species now called Loris tardigradus (the red slender loris). In 1758, he listed three species in the genus Lemur: Lemur tardigradus, Lemur catta (the ring-tailed lemur), and Lemur volans (now called Cynocephalus volans, the Philippine colugo). In 1911, Oldfield Thomas made Lemur catta the official species for the genus. In 1929, the International Commission on Zoological Nomenclature (ICZN) confirmed this decision.
The ring-tailed lemur shares similarities with ruffed lemurs (genus Varecia) and true lemurs (genus Eulemur). Its skeleton is very similar to that of true lemurs. In the past, all three groups were grouped together in the genus Lemur and later classified as the subfamily Lemurinae. In 1962, ruffed lemurs were moved to the genus Varecia. In 1988, scientists moved true lemurs to the genus Eulemur based on genetic and scent gland evidence. In 1991, some scientists suggested returning Eulemur and Varecia to the genus Lemur, but this idea was not widely accepted. The genus Lemur now contains only the ring-tailed lemur. Because genetic differences between the ring-tailed lemur and bamboo lemurs are very small, some scientists suggest merging all three groups.
Scientists disagree about how to classify lemurs in the family Lemuridae. Most primatologists support the current classification, but some continue to debate it.
In 1996, researchers Steven Goodman and Olivier Langrand noted that ring-tailed lemurs in the Andringitra Massif region have thicker fur, lighter color, and different tail patterns. In 2001, Colin Groves concluded that these differences do not represent a new subspecies. Later studies confirmed that these traits are normal variations, not new species. The thicker fur was likely an adaptation to cold temperatures, and lighter fur may result from more sunlight exposure. Genetic studies in 2000 also showed that the Andringitra population is not significantly different from other ring-tailed lemur populations.
Anatomy and physiology
The ring-tailed lemur is a relatively large lemur. Its average weight is 2.2 kilograms (4.9 pounds). Its head–body length ranges between 39 and 46 centimeters (15 and 18 inches), its tail length is 56 to 63 centimeters (22 to 25 inches), and its total length is 95 to 110 centimeters (37 to 43 inches). Other measurements include a hind foot length of 102 to 113 millimeters (4.0 to 4.4 inches), ear length of 40 to 48 millimeters (1.6 to 1.9 inches), and cranium length of 78 to 88 millimeters (3.1 to 3.5 inches).
The species has a slender frame and narrow face with a fox-like muzzle. The ring-tailed lemur's trademark is its long, bushy tail, which is ringed in alternating black and white bands. There are 12 or 13 white rings and 13 or 14 black rings, always ending in a black tip. The total number of rings is nearly equal to the number of caudal vertebrae (~25). Its tail is longer than its body and is not prehensile. Instead, it is used for balance, communication, and group cohesion.
The fur is very dense and can clog electric clippers. The chest and throat are white or cream. The back fur varies from gray to rosy-brown, sometimes with a brown patch around the tail, where the fur turns pale gray or grayish brown. The back is slightly darker around the neck and crown. The throat, cheeks, and ears have white or off-white fur that is less dense, allowing dark skin underneath to show. The muzzle is dark grayish, the nose is black, and the eyes are surrounded by black triangular patches. Facial vibrissae (whiskers) are present above the lips, on the cheeks, and on the eyebrow. Vibrissae are also found slightly above the wrist on the underside of the forearm. The ears are large compared to other lemurs and are covered in hair, which has only small tufts if any. While facial patterns may vary slightly between individuals, there are no obvious differences between the sexes.
Unlike most diurnal primates, but like all strepsirrhine primates, the ring-tailed lemur has a tapetum lucidum, or reflective layer behind the retina of the eye, that improves night vision. This layer is highly visible because the pigmentation of the eye’s back surface is spotty. The ring-tailed lemur also has a small foveal depression on the retina. Another shared trait with strepsirrhine primates is the rhinarium, a moist, glandular nose supported by the upper jaw and extending beyond the chin. The rhinarium divides the upper lip, which is attached to the premaxilla, preventing the lip from protruding and requiring the lemur to lap water instead of using suction.
The skin of the ring-tailed lemur is dark gray or black, even in areas where the fur is white. It is exposed on the nose, palms, soles, eyelids, lips, and genitalia. The skin is smooth, but the hands and feet have a leathery texture that helps with movement on the ground. The anus, located at the tail’s joint, is covered when the tail is lowered. The area around the anus and the perineum are covered in fur. In males, the scrotum lacks fur, has small, horny spines, and its two sacs are divided. The penis is nearly cylindrical, covered in small spines, and has two pairs of larger spines on both sides. Males have a small baculum (penis bone) compared to their size. The scrotum, penis, and prepuce are usually coated with a foul-smelling secretion. Females have a vulva with a thick, elongated clitoris that protrudes from the labia. The urethra’s opening is closer to the clitoris than the vagina, forming a "drip tip."
Females have two pairs of mammary glands (four nipples), but only one pair is functional. The anterior pair is close to the armpits. Both males and females have furless scent glands. Both sexes have small, dark forearm glands measuring 1 centimeter long, located on the inner surface of the forearm nearly 25 centimeters (9.8 inches) above the wrist. These glands are soft, compressible, have fine dermal ridges, and are connected to the palm by a hairless strip. Only males have a horny spur that covers this gland. The spur develops with age through secretions from an underlying gland with many tiny ducts. Males also have brachial glands on their shoulders near the armpit. These glands are larger than forearm glands, have short hair around the edges, and have a crescent-shaped opening. They secrete a foul-smelling, sticky brown substance. Brachial glands are rarely developed in females. Both sexes have apocrine and sebaceous glands in their genital or perianal regions, which are covered in fur.
The ring-tailed lemur’s fingers are slender, padded, mostly not webbed, and semi-dexterous with flat, human-like nails. The thumb is short and widely separated from the other fingers. Though set at a right angle to the palm, the thumb is not opposable because the joint is fixed. Like all strepsirrhines, the hand is ectaxonic (the axis passes through the fourth digit) rather than mesaxonic (the axis passes through the third digit) as seen in monkeys and apes. The fourth digit is the longest, slightly longer than the second digit. The fifth digit is also slightly longer than the second. The palms are long and leathery, with dermal ridges to improve grip. The feet are more specialized than the hands. The big toe is opposable and smaller than the big toes of other lemurs. The second toe is short, has a small pad, and has a toilet-claw (grooming claw) for grooming fur that the mouth cannot reach. This trait is shared among nearly all living strepsirrhine primates. Unlike other lemurs, the ring-tailed lemur’s heel is not covered in fur.
The ring-tailed lemur has a dentition of 2.1.3.3 2.1.3.3 × 2 = 36, meaning each side of the jaw has two incisors, one canine, three premolars, and three molars. Its deciduous dentition is 2.1.3 2.1.3 × 2 = 24. Permanent teeth erupt in this order: m1/1 (first molars), i2/2 (first incisors), i3/3 (second inc
Ecology
The ring-tailed lemur is active during the day and partly on the ground. It is the most ground-dwelling lemur species, spending about 33% of its time on the ground. However, it still spends time in trees, with 23% of its time in the mid-level canopy, 25% in the upper-level canopy, 6% in the highest layer of the forest, and 13% in small bushes. When traveling, 70% of the time is spent on the ground.
Troop size, home range, and population density depend on the area and the availability of food. Troops usually have 6 to 25 members, though some have more than 30. The average troop has 13 to 15 members. Home ranges vary between 6 and 35 hectares (15 to 86 acres). Ring-tailed lemurs protect their territory, but their ranges often overlap. When conflicts happen, they are hostile. A troop typically stays in one area for three to four days before moving. When they move, they travel about 1 kilometer (0.62 miles) on average. Population density ranges from 100 individuals per 1 square kilometer (0.39 square miles) in dry forests to 250–600 individuals per square kilometer in gallery and secondary forests.
The ring-tailed lemur has both natural and introduced predators. Natural predators include the fossa (Cryptoprocta ferox), the Madagascar harrier-hawk (Polyboroides radiatus), the Madagascar buzzard (Buteo brachypterus), and the Madagascar ground boa (Acrantophis madagascariensis). Introduced predators include the small Indian civet (Viverricula indica), domestic cats, and domestic dogs. Other predators are black kites and Malagasy giant hognose snakes.
The ring-tailed lemur is found in southern and southwestern Madagascar and lives in higher areas than other lemurs. It lives in deciduous forests, dry scrub, montane humid forests, and gallery forests (forests along rivers). It prefers gallery forests, but many of these have been cleared for farming. Temperatures in its range can vary from −12°C (10°F) in the Andringitra Massif to 48°C (118°F) in the spiny forests of Beza Mahafaly Special Reserve.
This species is found as far east as Tôlanaro, inland toward the Andringitra mountains, in the spiny forests of southern Madagascar, and north along the west coast to the town of Belo sur Mer. Historically, its range in the west extended to the Morondava River near Morondava. It is still found in Kirindy Mitea National Park, though in very low numbers, but not in Kirindy Forest Reserve, which is north of Morondava. Its distribution is uneven, with population numbers varying widely across its range.
The ring-tailed lemur can be seen in five national parks in Madagascar: Andohahela National Park, Andringitra National Park, Isalo National Park, Tsimanampetsotse National Park, and Zombitse-Vohibasia National Park. It is also found in Beza-Mahafaly Special Reserve, Kalambatritra Special Reserve, Pic d'Ivohibe Special Reserve, Amboasary Sud, Berenty Private Reserve, Anja Community Reserve, and marginally in Kirindy Mitea National Park. It lives in unprotected forests such as Ankoba, Ankodida, Anjatsikolo, Anbatotsilongolongo, Mahazoarivo, Masiabiby, and Mikea.
In protected areas, the ring-tailed lemur lives with up to 24 other lemur species, covering all living genera except Allocebus, Indri, and Varecia. Historically, it lived with the critically endangered southern black-and-white ruffed lemur (Varecia variegata editorum) in Andringitra National Park, but no recent sightings of the ruffed lemur have been reported.
In western Madagascar, ring-tailed lemurs and red-fronted lemurs (Eulemur rufifrons) live in the same area but interact little. Their diets overlap, but the ring-tailed lemur eats more varied foods and spends more time on the ground.
The ring-tailed lemur is an opportunistic omnivore that mainly eats fruits and leaves, especially from the tamarind tree (Tamarindus indica), known locally as kily. Tamarind can make up as much as 50% of its diet, especially during the dry season. It eats from up to 36 different plant species, including flowers, herbs, bark, and sap. It has been seen eating decayed wood, soil, spider webs, insect cocoons, arthropods (spiders, caterpillars, cicadas, and grasshoppers), and small vertebrates (birds and chameleons). During the dry season, it eats more varied foods.
Behavior
Ring-tailed lemurs live in groups called "troops." These groups have many males, and the core of each group is a family line led by females. Female lemurs are socially dominant over males in all situations, including when they eat. They use actions like lunging, chasing, grabbing, and biting to show dominance. Young females do not always take the same rank as their mothers, and young males leave the troop between three and five years old. Both males and females have separate rankings. Female rankings are clear, while male rankings depend on their age. Each troop has one to three high-ranking adult males who spend more time with females and lead the group. Males who are new to the group, older, or young and still with their birth group are often lower in rank. These males usually stay on the edges of the group and are less involved in activities.
For males, the social structure can change with the seasons. Between December and May, some males move between groups. Established males move on average every 3.5 years, while younger males may move about every 1.4 years. When groups become too large and food is scarce, the group may split into smaller groups.
In the morning, ring-tailed lemurs sit in the sun to warm themselves. They often sit in a posture called the "sun-worshipping" position, with legs stretched out, not crossed. They may use nearby branches for support. Sunning is often done in groups, especially during cold mornings. At night, the troop splits into smaller groups that huddle together to stay warm.
Although ring-tailed lemurs walk on four legs, they can stand on their hind legs for displays of aggression. When threatened, they may jump and use their short nails and sharp teeth in a behavior called "jump fighting." This is rare except during the breeding season when competition for mates is high. Other aggressive behaviors include staring to intimidate others and showing their lips in a submissive way.
Disputes with other troops happen sometimes, and the dominant female is responsible for protecting the group's area. Conflicts may involve staring, chasing, or physical fights, and usually end with the group retreating to the center of their territory.
Smell is very important for communication in ring-tailed lemurs. Both males and females mark their territory using scent glands on their body. To mark vertical surfaces, they stand on their hands and feet, using their feet to grip the highest point while applying their scent. Males use glands on their arms and wrists to mark territory and show dominance. They scrape the spiny bumps on their wrists against trees to spread their scent, a behavior called "spur-marking."
During aggressive displays, males use a behavior called "stink fighting." They spread scent from their arm and wrist glands onto their tails and wave the scented tail at rivals.
Ring-tailed lemurs also mark their territory with urine. Regular urination involves raising the tail slightly and releasing a stream of urine. Urine marking is different: the tail is held high, and only a few drops are used. This behavior is mainly done by females near the edges of their territory and during the mating season. It may help communicate about reproduction between groups.
Ring-tailed lemurs are very vocal and use many sounds to keep the group together and warn of danger. Simple calls, like a "purr," show happiness. Complex calls, such as clicking sounds and yaps, are used when the group is trying to scare away predators. Other common calls include "moans" and "wails," which help the group stay connected.
Ring-tailed lemurs are polygynandrous, meaning many females mate with many males. The dominant male in the troop usually mates with more females than other males. Mating often happens during the breeding season, which runs from mid-April to mid-May. A female may signal her readiness to mate by showing her backside and looking at a male over her shoulder. Males check the female's genital area to see if she is ready. While most mating happens within the troop, females may also mate with males from other groups.
Females are ready to mate for 4 to 6 hours each day during the breeding season. They spread out their readiness so that each female has a different day to mate, reducing competition. Mothers nurse their young during the wet season, when food is plentiful, and give birth during the dry season, when food is less available. A pregnancy lasts about 135 days, and babies are born in September or October. Usually, one baby is born, though twins can happen. Newborns weigh about 70 grams and are carried on the mother's chest for the first two weeks, then on her back.
Young lemurs start eating solid food after two months and stop nursing after five months. They reach sexual maturity between 2.5 and 3 years old. Males are not usually involved in raising babies, but all members of the troop, regardless of age or gender, may help care for the young. Some females have been seen caring for babies that are not their own, and males sometimes harm babies from other groups. Due to harsh conditions, many babies die before their first birthday. Only about 30% of babies survive to adulthood. The oldest known wild female lived to be 20 years old, while the oldest male lived to be 15. In captivity, some ring-tailed lemurs have lived up to 27 years.
Historically, studies on learning and intelligence in non-human primates focused on monkeys and apes, while lemurs and similar animals were often ignored or considered less intelligent. This may be because early experiments were designed for simians, making them unsuitable for lemurs. For example, simians are better at tasks that require certain behaviors, which lemurs may not naturally do.
Conservation status
The ring-tailed lemur is listed as endangered by the IUCN since 2014 and has been protected under CITES Appendix I since 1977. This means it is illegal to trade wild-caught ring-tailed lemurs. While other lemur species are also endangered, the ring-tailed lemur is considered a flagship species because it is easily recognized. As of 2017, only about 2,000 ring-tailed lemurs are estimated to live in the wild, which makes their risk of extinction more serious than previously thought.
Three main threats affect ring-tailed lemurs. First, habitat destruction has occurred for thousands of years. Humans began clearing forests for farming and grazing land long ago. Hardwood trees have been cut for fuel and building materials, and mining and overgrazing have also damaged the environment. Today, about 90% of Madagascar’s original forests are gone. Growing populations in the southwest have increased the need for fuel wood, charcoal, and lumber. Fires from clearing grasslands and slash-and-burn farming also harm forests. A second threat is people hunting lemurs for food, fur, or as pets. A third threat is droughts in southern Madagascar, which can harm already declining populations. For example, a severe drought in 1991 and 1992 caused many young lemurs and females to die at the Beza Mahafaly Special Reserve. The population dropped by 31% and took nearly four years to recover.
Ring-tailed lemurs live in several protected areas, each with different levels of protection. At the Beza Mahafaly Special Reserve, conservation efforts include involving local people and students in research and management. Livestock management is used near the reserve’s edges, and ecotourism helps support nearby communities.
Although their wild habitat is shrinking, ring-tailed lemurs reproduce easily and do well in captivity. This, along with their popularity, makes them the most common lemur in zoos worldwide. More than 2,500 live in captivity as of 2009, and they are also the most common captive primates. Conservation groups, such as the Duke Lemur Center in North Carolina, the Lemur Conservation Foundation in Florida, and the Madagascar Fauna Group at the Saint Louis Zoo, work to protect them. Because captive breeding has been successful, reintroducing lemurs to the wild might be possible if wild populations decline. Experimental releases on St. Catherines Island in Georgia showed that captive lemurs can adapt to the wild and behave naturally, but reintroduction is not currently happening.
Droughts can also be managed through efforts like watering troughs and planting fruit trees, as seen at the Berenty Private Reserve in Madagascar. However, these actions sometimes cause concern because they may interfere with natural population changes. Scientists believe the ring-tailed lemur’s ability to reproduce quickly evolved as a way to survive in its challenging environment.
Cultural references
The ring-tailed lemur is called maky (pronounced [ˈmakʲi̥]) or hira (pronounced [ˈhirə] or [ˈir]) in the Malagasy language. It is the most well-known native primate in Madagascar and serves as the symbol for Madagascar National Parks (formerly ANGAP). A clothing brand named Maki was created in Madagascar and now sells items across the Indian Ocean islands. The brand is named after the lemur because of its popularity, even though its logo shows a sifaka and uses the French spelling maki.
The first written record of the ring-tailed lemur in Western sources appeared in 1625. English traveler Samuel Purchas described the animal as similar in size to a monkey and noted its long, black-and-white ringed tail like a fox’s. In 1788, Charles Catton included the species in his book Animals Drawn from Nature and Engraved in Aqua-tinta, referring to it as the "Maucauco" and classifying it as a type of monkey.
The species gained more attention through the Animal Planet series Lemur Street and the character King Julien from the Madagascar films and TV shows. A 1996 Nature documentary titled A Lemur's Tale focused on a group of ring-tailed lemurs at the Berenty Reserve. The group included an infant named Sapphire, who had nearly white fur, bright blue eyes, and the typical ringed tail.
A ring-tailed lemur appeared in the 1997 comedy film Fierce Creatures, starring John Cleese. Cleese later hosted the 1998 BBC documentary In the Wild: Operation Lemur with John Cleese, which followed efforts to reintroduce black-and-white ruffed lemurs into the Betampona Reserve. Part of the project’s funding came from Cleese’s donation of money earned from the London premiere of Fierce Creatures.