Lemurs

(Lemuridae)

Class Mammalia

Order Primates or Primata

Family Lemuridae

Thumbnail description
Arboreal primates with monkeylike bodies and foxlike heads

Size
Approximately that of a house cat; adult head-and-body length 11–22 in (28–56 cm), tail length 11–24 in (28–60 cm), adult weight 4.4–10 lb (2–4.5 kg)

Number of genera, species
2 genera; 9 species

Habitat
Tropical and subtropical forests

Conservation status
Critically Endangered: 2 species; Endangered: 1 species; Vulnerable: 5 species

Distribution
The island of Madagascar and the Comoro Islands

Evolution and systematics

The evolutionary history of the Malagasy prosimians has been, until recently, one of the knottiest in the life sciences. As of 2000, cladistic analyses, genetic and mitochondrial DNA studies, and morphological comparisons support a monophyletic (single species) origin of all Malgasy prosimians from a founder species that rafted on vegetation from Africa to Madagascar in the early Eocene epoch (55 to 34 million years ago).

There are at least 60 known species of Malagasy prosimians, living and recently extinct, all generally lumped under the umbrella name "lemurs." The total includes 15 large-bodied species, most with unique adaptations, that became extinct within the last 200–300 years. "Lemuridae" is used here as referring to the so-called true lemurs, house cat-sized with fairly long, fox-like muzzles.

Lemurs are prosimians (suborder Prosimii), the term "prosimian" is loosely translated as "pre-monkey" and covers several lines of primate evolution diverging from those of the anthropoids (suborder Anthropoidea: monkeys, apes, and hominids). Living prosimian species include the prosimians of Madagascar (superfamily Lemuroidea), the lorisoids, and the tarsiers. The lorisoids (superfamily Lorisoidea) include the galagos (bushbabies) and pottos of the African mainland and the lorises of Southeast Asia. There are only three living species of tarsiers (infraorder Tarsiiformes, superfamily Tarsiodea), small, headlight-eyed, goblinesque arboreal primates found on some of the Southeast Asian islands.

The eye socket of the skull is open in prosimians but closed in anthropoids. Prosimians have mostly nails on their digits, except for the second digit of the hind foot, which carries a claw or clawlike nail used for self-grooming; anthropoids have only nails on all digits (with the distant exception of the neotropical marmosets and tamarins). Prosimians' lower canines and incisors are modified into a comblike structure used as a grooming tool; anthropoids have no such structure.

Suborder Strepsirrhini ("wet nose") covers lemurs and lorisoids, since those species keep the generalized mammalian condition of noses wet by self-licking to facilitate the olfactory sense, obvious in animals like dogs and cats. In strepsirrhines, the upper lip is divided, again as in dogs and cats, to

make way for the frequent nose-lapping tongue. Tarsiers, monkeys, apes, and hominids are placed in suborder Haplorrhini ("dry nose"), since they have discarded the split lip and the wet nose, coming to rely more on vision and less on olfaction.

There are several differing classifications of lemur species, all in fluctuation as the latest studies in morphology and DNA comparison studies of lemurs reveal new interrelationships among species. The other families of living Malagasy prosimians, not covered in this entry, are Cheirogaleidae, the dwarf and mouse lemurs; Lepilemuridae, the weasel or sportive lemurs; Indriidae, covering the indris, sifakas, and avahis; and Daubentoniidae, the specialized, enigmatic aye-aye.

Physical characteristics

Lemuridae are arboreal primates, the size of house cats, with bodies, limbs, hands and feet much like those of monkeys, somewhat foxlike heads with long muzzles, and large, brightly hued, round, owl-like eyes. Adult Lemuridae head-and-body length ranges 11–22 in (28–56 cm) and tail length 11–25.5 in (28–65 cm). The tail length in most species is longer than the head-and-body length. Adult weights run 4.4–10 lb (2–4.5 kg). Bodies and limbs are gracile, the hind limbs longer than the forelimbs. The pelage is dense, soft, woolly or cottony, and rather long. Species may carry face or neck ruffs of long fur. Coat colors and patterns vary considerably among species. Some species are sexually dichromatic.

The eyes are set close for binocular vision, and in most species are brilliantly colored. Olfactory communication being important, lemuridae are equipped with scent glands in various parts of their bodies, the exact number and location

varying throughout genera and species. The animals rub exhudations from the glands on various spots and objects to mark territory.

All lemuridae (and all prosimians) carry grooming combs, modified lower incisors and canines that form a comblike structure that the animals use for grooming themselves and others of their group. Mutual grooming is a social glue, maintaining and reinforcing bonds among the members of a group.

The ears are partially furred and not especially prominent, except in Lemur catta, where they are large and triangular, resembling the ears of housecats.

The hands and feet are more or less monkeylike. The pollex (thumb) and hallux (big toe) are set off more or less at right angles to the other digits and are opposable, ensuring firm holds on tree branches. The palms and soles are deeply ridged for a firm grip. A claw or clawlike nail, used for self-grooming, is present on the second toe of each hind foot.

Distribution

Madagascar and the Comoro Islands.

Habitat

Lemur species have adapted to several varied habitats within their range, including humid lowland and montane

tropical forests, dry scrub, dense gallery forest, dry tropical deciduous forest, sparse rocky areas, and occasionally grassland.

Behavior

All Lemuridae species are arboreal, although most spend some minimum time on the ground. The ringtailed lemurs are notable for spending about half their foraging time on the ground and can live in treeless areas.

In the trees, Lemuridae walk and run quadrupedally along the tops of tree limbs and leap between trees. At rest, they sit upright or lay down. The tail is about as long as the body, thickly furred, and used for balance and for steering during jumps.

The Lemuridae are for the most part diurnal foragers, with some exceptions. Lemur mongoz alternates between diurnal and nocturnal activity in response to season and food availability.

All the Lemuridae species are social, but the exact formalities vary among species. The number of individuals within a group may range anywhere from two to 20. There may be large groups that break up into smaller foraging groups during the day, then reconglomerate at nightfall into the original,

single, large group. There may be small family groups of permanently bonded males and females and their offspring.

Groups maintain their cohesion by means of the frequent and all-important activity of mutual grooming with the "grooming comb" derived from the lower incisor and canine teeth.

Lemuridae societies are female-dominant. Females have priority in choosing mating partners and helping themselves to larger amounts of food. A single female leads a typical group of females and males in foraging and sheltering. Either sex has its own dominance hierarchy.

Lemuridae are territorial. Abutting same-species territories may or may not overlap. When neighboring foraging troops meet at territory boundaries, both react by staging hyperactive bouts of alarm calls and branch-shaking.

The Lemuridae have an almost musical range of vocalizations for various needs. There are calls for greeting, territorial assertions, contact and threats between in-group or out-group individuals, and alarm calls that vary according to the type of threat.

Feeding ecology and diet

Diet is herbivorous overall, with some omnivory. Plant foods include flowers, pollen, nectar, fruits, leaves, seeds, and seed pods. Less often on the menu are insects and other invertebrates, small vertebrates, and birds' eggs.

Reproductive biology

All species of Lemuridae mate from April through June, the females have a gestation period of about 4.5 months and give birth from August through October, generally coinciding with the beginning of the monsoon season. When plant growth resumes, animals wake from torpor and new food becomes available.

The females of all the Lemuridae genera except Varecia have but one pair of mammae, while Varecia carries six pairs. A female reaches sexual maturity at two years old and usually starts bearing young, annually, in her third year. There are generally one or two young per birth, although

Varecia may have up to six young (and six mammae to feed them all).

For the first four weeks of life a newborn young rides beneath the mother's body, gripping the ventral fur and hugging the torso. After the fourth week, the youngster switches to riding on the mother's back. At about one month of age it starts wandering and exploring on its own. In two months, it begins sampling solid food, and until weaning—at about five to six months—will gradually replace its milk diet with solid food. The exact timing of these different stages of development may vary among species.

Lemurs in the wild can live perhaps 20 years. The record for longevity in captivity among Lemuridae is 39 years for a hybrid of Lemur macaco and Lemur fulvus.

Conservation status

Since the Lemuridae are primarily arboreal, rampant destruction of Madagascar's remaining forests for the purpose of agriculture and grazing puts all the lemuridae species at risk. Taking a lesser but significant toll is the hunting and trapping of lemurs for both food and for the illicit market in exotic pets.

Significance to humans

Lemuridae species are hunted and trapped for food. Some are kept as pets or shipped abroad for the exotic pet trade. A few species are blamed for raiding crops and are consequently hunted and trapped.

On the brighter side, ecotourism has taken off in Madagascar, and lemurs in the wild and their habitats have become a substantial tourist draw, bringing in valuable foreign exchange to Madagascar, one of the world's poorest countries. Through the scientific study of lemurs we can learn more about adaptive evolution and speciation. Lemurs have become rallying symbols for conservation because they are beautiful, charming, and fascinating animals.

Species accounts

List of Species

Ringtailed lemur

Lemur catta

taxonomy

Lemur catta Linnaeus, 1758, Madagascar.

other common names

French: Maque, maki mococo, maki; German: Katta; Spanish: Lemur colianillado.

physical characteristics

Adult head-and-body length of 15–18 in (39–46 cm), tail length of 22–24 in (56–62 cm), and adult body weight of 6.5–7.75 lb (3–3.5 kg). People without foreknowledge of lemurs, on seeing ringtailed lemurs in zoos or pictures, may regard them as some sort of aberrant raccoon, suggested by the overall shape and especially by the bandit-masked faces and vividly striped tails.

The pelage is dense and soft. The main body color is light gray to gray-brown on the flanks, rump, and limbs, light reddish-brown along the back, and dark gray on the crown and back of the neck. The head is the least monkey-like feature, with its long, foxlike muzzle. The triangular ears are covered with white fur. The forehead, bridge of the snout and proximal half of the muzzle and face are white. The distal half of the muzzle is dark gray or black. The eyes, each encircled by a prominent black ring, are bright red-brown or orange, with less of the "dead fish" stare common among lemur species. The most arresting feature is the tail, as long as the head and

torso, and emphatically striped bright white and jet-black, with 13 or 14 black rings. Hindlimbs considerably longer than the forelimbs, giving the animals a somewhat hunched, leaning-forward look as they stand on all fours or locomote.

Individuals are equipped with scent glands on wrists (carpal or antebrachial glands), arms (brachial glands), and chests with which they mark territory and foraging routes with exhudations. Males' wrist glands are further gifted with small, horny, thornlike outcrops with which the males gouge scars into tree trunks and branches to add a visual component to their scent markings.

distribution

Ringtailed lemurs live throughout southern Madagascar, from Tôlanaro (Fort-Dauphin) on the east coast and as far north as Morandava on the west coast, with a separate population in Andringita Natioanl Park in south-central Madagascar.

habitat

Ringtails are comfortable in several types of indigenous forest, from dry scrub forest to dense, closed-canopy gallery (river-side) forest. They also reside in (or at least take excursions into) indigenous, dry-adapted spiny forests, which are extensive in the south of Madagascar. A separate population has taken to living in dry, rocky, treeless areas in Andringitra National Park in south-central Madagascar, perfectly at home on rocky outcrops and vertical cliffs. These ringtails differ from the general run in having darker pelts and fewer rings on their tails. They have created and colonized their own unique ecological niche and are the only living lemur species to have adapted to a treeless environment in the wild.

behavior

Lemur catta is among the most adaptable lemur species and the one that spends the most time on the ground, although they are just as comfortable in the trees. Ringtails live in groups of 5–25 individuals, an average being 14. Larger groups form a core group of adult females and infants, juveniles, and one or more high-ranking males. Females dominate males, thereby getting first pick of food and mating partners, but there is not always a single, individual leader for the entire group. A female remains within the group in which she was born, while males tend to wander among groups.

A ringtailed lemur group forages in a range of 15–22 acres (6–9 ha) in densely forested areas and up to 57 acres (23 ha) in scrub. Ringtailed lemur territories border on one another without overlap.

Ringtailed lemurs are diurnal, starting the day with a "sun-worship" posture, sitting upright on the ground, arms held out from the sides and resting on the knees, palms open. This gesture is seen in other lemur species and serves to soak up sunlight and thus warm the body in the cool mornings. As the day warms, the troop goes searching for food and forages until noon, when the group naps in trees or on the ground during the hottest part of the day. In the afternoon, they rouse and forage again before nightfall, at which time they take to the trees for sleeping.

In-group disputes are common and may culminate in "stink-fights" between rival males. One male positions his wrist glands close together and drags his tail between them, coating the tail fur with his scent. Then he flicks and waves the tail at the rival, who may respond in kind or back off.

Ringtailed lemurs have a range of identification and alarm calls. The most often-heard sounds are a very cat-like meow for group cohesion, though the sound is a little more high-pitched and songlike. The animals also make rapid, high-pitched yapping or barking sounds for threat or alarm.

Although arboreal animals, ringtail lemurs walk and run quite comfortably and efficiently on the ground. When a ring-tail troop travels on the ground, the members keep their tails raised straight into the air, like flags, for group cohesion.

feeding ecology and diet

The ringtailed lemur diet is varied but primarily vegetable, including fruit, leaves, flowers, herbs, tree sap, bark, and other plant parts, although they may at times supplement their diets with insects and small vertebrates. A favorite fruit, when in season, are the seed pods of the tamarind tree (Tamarindus indica). The only edible parts are the sticky, sweet-tart arils, or coatings on the seeds.

reproductive biology

Females reach sexual maturity at two years of age, birth their first young at three years and follow with annual births. Males reach sexual maturity at two and a half years old, but must contend with older, dominant males who may curtail any mating moves on the part of young males.

Mating begins in mid-April. During the mating season, females enter estrous for only a few hours of a single day, although all the females of a group will enter estrous within two weeks of one another. The males, driven by hormones, fight madly among themselves for mating privileges during that narrow time window. Females typically mate with more than one male, often with one of the males being from another troop. The young are born in August and September. One or sometimes two are born, depending on food abundance.

The newborn at first clings to the fur of its mother's under-side, but in three days it begins moving about on its mother's body, still grasping the pelt. By two weeks of age, the youngster is riding stomach-down on the mother's back. By two and a half months it leaves the mother to play with other young, explore and sample solid foods, though it is still carried by the mother whenever the group moves. The youngster spends more time per day on its own, eating more solid food and taking less milk until final weaning at 5–6 months of age.

Females in a group with newborns show considerable "aunt behavior," handing infants about, even nursing other females' infants, and attending groups of young as they play.

conservation status

Vulnerable. Multiple threats include deforestation and hunting for food and the illegal pet trade.

significance to humans

Ringtailed lemurs are readily visible in several protected areas in southern Madagascar and have thereby become a banner species for ecotourism. In Berenty Reserve, near Fort Dauphin, at least one troop of ringtails has become so tame that visitors can follow the group about its business throughout the day. Ringtailed lemurs have become reliable ecotourism magnets and thereby bring visitors, cash, and business into Madagascar.

Mongoose lemur

Lemur mongoz

taxonomy

Lemur mongoz Linnaeus, 1766, Anjouan Island, Comoros.

other common names

French: Lémur mongoz; German: Mongozmaki; Spanish: Lemur mangosta.

physical characteristics

Adults have an average head-and-body length of 14 in (35 cm), tail length of 19 in (48 cm), and adult body weight of 4.5 lb (2 kg). Pelages are colored gray on the heads, forelimbs, and shoulders, dark grey on the back, accentuated by a reddish brown beard. Often the muzzle is white. The eyes are a lustrous red-brown.

distribution

Mongoose lemurs are found in forests in northwestern Madagascar and on the islands of Moili (Moheli) and Ndzouani (Anjouan) of the Comoros. In northwestern Madagascar, the range of mongoose lemurs extends from the Bay of Narinda in the north to as far south as the Betsiboka River.

The Comoros are a volcanic-origin island group in the Mozambique Channel, northwest of Madagascar, midway between Madagascar and mainland Africa. The mongoose lemur is one of only two lemur species living naturally outside of mainland Madagascar, the other being the brown lemur (L. fulvus), also living on the comoro. Both species are probably feral descendants of individuals brought by man from Madagascar to the islands.

habitat

Mongoose lemurs live in tropical dry deciduous forests on Madagascar and in humid lowland and montane tropical forests on Moheli and Anjouan.

behavior

Mongoose lemurs live in groups of three or four individuals, a female-male pair and their pre-weaned young, the adult female dominant to the adult male. Territories of neighboring groups often overlap. Groups encountering each other at the edges or overlap zones of abutting territories respond with elaborate, noisy intimidation displays. Yet, groups of mongoose lemur and brown lemur may forage alongside one another, or even intermingle as they forage.

Mongoose lemurs on mainland Madagascar forage diurnally or nocturnally, depending on the season, a prime changeover time from day to night activity taking place at the beginning of the dry season, when food becomes scarce. Nocturnal feeding reduces competition from other species.

feeding ecology and diet

Mongoose lemurs help themselves to flowers, nectar, pollen, fruits, and leaves.

reproductive biology

Monogamous. Young are born in mid-October, usually one per litter, with annual births. Male mongoose lemurs are born with white beards that change to reddish-brown at five to six weeks of age. Newborn young cling to their mothers' undersides for the first three weeks. At five weeks, youngsters begin to walk independently and sample solid food. Nursing continues, with the mother's milk gradually replaced by solid food, until weaning at 5–6 months of age. Young mongoose lemurs reach sexual maturity by two or three years, at which time male and female young are driven from the parental group. Intergroup encounters are rare, however, and often result in agitation, vocalizations, and scent marking.

conservation status

Vulnerable. Mongoose lemurs exist in only a few limited populations threatened by deforestation. The species is hunted for food and for the illegal pet trade.

significance to humans

Mongoose lemurs are hunted for food.

Black lemur

Lemur macaco

taxonomy

Lemur macaco Linnaeus, 1766, Madagascar.

other common names

English: Blue-eyed lemur, Sclater's lemur; French: Lémur macaco, maki dimorphisme; German: Mohrenmaki; Spanish: Lemur negro.

physical characteristics

There are two subspecies, black lemurs or Lemur macaco macaco and blue-eyed lemurs or Sclater's lemurs, Lemur macaco flavifrons. Blue-eyed lemurs are the only primates besides humanity to bear blue eyes. The trait is probably due to the genetics of an isolated population that enabled recessive genes to express.

Matings of black and blue-eyed lemurs produce only brown-eyed young.

The two subspecies are nearly identical in shape, size, and behavior, but they live in different habitats and show different colors in pelage and eyes. An adult individual of either sub-species has an average adult head-and-body length of 16 in (41 cm), tail length of 22 in (55 cm), and an adult body weight of 5 lb (2.4 kg).

Both subspecies are sexually dichromatic. Lemur macaco macaco males have jet-black fur with brown highlights from nose tip to tail tip, while female pelages sport mixtures of brown, reddish brown, and gray on their backs, with off-white undersides and black faces. Both sexes bear somewhat raggedlooking ruffs of fur framing the sides of their faces, black in males and off-white in females. Males' eyes are dark brown; females' are golden to orange. Lemur macaco flavifrons males are completely black but with blue eyes. Females have a light reddish brown coat with a white brow-bar above the blue eyes.

distribution

Lemur macaco macaco are found in the northwest humid tropical forests of Madagascar, including those on the islands of Nosy Be and Nosy Komba off the northwest coast. On Nossi Komba, black lemurs have become part of the tourist circuit, semi-tame individuals obligingly cadging bananas from visitors and locals, and lounging about on rooftops. Lemur macaco flavifrons are found in northwestern Madagascar in a range south and separate from that of L. m. macaco, from the Andranomalaza River further southward to the Sandrakota River.

habitat

Black lemurs are at home in humid and dry primary and secondary tropical forest, and humanity-altered land such as timber plantations and combinations of partially cleared forests and plantings of coffee, citrus, and cashew.

behavior

Black lemurs are diurnal and social, groups being made up of 2–15 individuals, often with more males than females. Small groups may forage within their own territories during the day, but several such groups may bunch together for the night.

feeding ecology and diet

Ripe fruit, leaves, flowers, and insects.

reproductive biology

Polygamous. Black lemurs reach sexual maturity at two years. Mating in both subspecies takes place from June through July. Mothers give birth to one or two young after a gestation period of about 126 days.

conservation status

Vulnerable. Deforestation and hunting are the primary threats.

significance to humans

Because of the large amounts of fruit in their diet, black lemurs may indirectly benefit man by being important dispersers of fruit seeds in forests, thus contributing to the overall health of those forests.

Variegated lemur

Varecia variegata

taxonomy

Lemur variegata Kerr, 1792, Madagascar.

other common names

English: Ruffed lemur.

physical characteristics

Varecia variegata is the largest in body size of the living Lemuridae species. The adult head-and-body length is 20–22 in (51–56 cm), tail length is 22–25.5 in (56–65 cm), and adult body weight is 7–10 lb (3.2–4.5 kg). Females are larger than males. The ruffed lemur carries a scent gland on its neck, and females have six mammae.

There are two subspecies: black and white ruffed lemurs, Varecia variegata variegata and red ruffed lemurs, Varecia variegata rubra. Black and white ruffed lemurs have a black tail, a body patched black and white in varied arrangements, and a white ruff around a black face. The fur is rather long, with a soft, cottony texture. The eyes are light gray or gray-green. Red ruffed lemurs have a black tail, face and hands, a reddish-brown body, and a white patch on the back of the neck and across the shoulders. The undersides and inner limbs are black. The eyes are golden and look almost self-luminous contrasted with the dark face. The ruff is like that on V. v. variegata but colored off-white.

distribution

Lowland and montane rainforest belt paralleling the east coast of Madagascar, including the forests of the Masoala Peninsula in northeastern Madagascar.

habitat

Black and white ruffed lemurs live throughout the montane rainforests paralleling the east coast of Madagascar, and on the small island of Nosy Mangabe in the Bay of Antongil. Red ruffed lemurs live only in the forests of the Masoala Peninsula, which forms the Bay of Antongil in northeastern Madagascar. The Masoala forests include some of the few large remnants of Malagasy east coast lowland tropical rainforest. Part of the forested peninsula is now Masoala National Park.

behavior

Black and white ruffed lemurs live in small groups averaging five individuals, a bonded female-male pair and offspring. Females are dominant to males. Larger, female-led groups form during the rainy season, when food is more abundant, and fall apart into smaller groups or lone individuals during the food-scarce dry season.

Ruffed lemurs are the most committedly arboreal of the living Lemuridae species, spending their days in the upper levels of their rainforest homes. An individual moves through this area by walking or running on all fours on larger branches and jumping between trees, but more slowly and carefully than smaller species of the Lemuridae. Ruffed lemurs are crepuscular, being most active during morning and evening twilight.

Ruffed lemurs sound off with an impressively varied repertoire of loud, strange calls, including roaring alarm calls. They can produce up to a dozen differing alarm calls at the sight of boas, raptors, and the fossa (Cryptoprocta ferox), a small but aggressive predatory mammal endemic to Madagascar.

feeding ecology and diet

Completely herbivorous and consists almost entirely of fruit, sometimes varied with leaves, nectar, and seeds.

reproductive biology

Polygamous. Females may bear litters of up to six young, something of an oddity among primates. They carry six mammae although the most common count is three young per litter. The gestation period lasts 90–102 days, shorter than is common for the Lemuridae.

The young do not hold onto or ride the mother as in other Lemuridae species; instead, the mother carries them about, one at a time, in her mouth, and builds a nest of leaves for holding them while she goes off seeking food on her own. The mother blankets the nest with her own fur, which she pulls from her pelt.

The young mature rapidly. They begin leaving the nest tentatively and temporarily at only three weeks of age. They are able to climb expertly in trees at five weeks and are as active and agile as their mother at seven weeks. Full weaning comes about at around five and a half months. At six months the young have nearly reached adult size.

conservation status

Endangered. Prospects for the future of ruffed lemurs is a mix of unfavorable and favorable. Black-and-white ruffed lemurs are severely threatened by the loss of rainforest habitat along the east coast of Madagascar, in the lowlands and in the mountains paralleling the coast. Nearly all the original eastern lowland rainforest is gone, converted into farmland, and the mountain rainforests are shrinking yearly as they are cleared. The Malagasy hunt and trap the animals for food. They also fall victim to a lively worldwide black market in exotic pets. Red-ruffed lemurs are somewhat better off in the partially protected forests of Masoala Peninsula, but deforestation and hunting continue to decrease their numbers.

significance to humans

Variegated lemurs are hunted for food and captured for the illegal exotic pet trade. However, captive-born ruffed lemurs have been reintroduced to the wild and serve as educational aids in Madagascar.

Brown lemur

Lemur fulvus

taxonomy

Lemur fulvus Geoffroy, 1796, Madagascar.

other common names

English: Fulvous lemur; French: Lémur brun; German: Brauenmaki.

physical characteristics

There are seven subspecies of brown lemur. The subspecies are similar in body size and behavior, but pelage color and markings vary among them, and all subspecies show sexual dichromatism. An adult brown lemur of any subspecies has a head-and-body length of 15–20 in (38–50 cm), tail length 18–24 in (47–60 cm). Adult body weight runs 4.5–9 lb (2.1–4.2 kg).

distribution

Brown lemurs are found throughout all of Madgascar's remaining dry western and central forests. In the west, they live all through the belt of forest starting north of the Betsiboka River, to the northwest corner of Madagascar. Subspecies Lemur fulvus mayottensis lives on the island of Mayotte in the Comores. Along with mongoose lemurs (L. mongoz), brown lemurs are the only lemurs living naturally outside of Madagascar.

Of the seven subspecies, the exact distributions of the six on mainland Madgascar have not been determined. Only L. f. mayottensis, in its limited range on Mayotte Island, has a confirmed distribution.

habitat

Brown lemurs live in tropical deciduous forest along the west coast of Madagascar, in scattered forest fragments on Madagascar's high plateau, and in lowland and montane rainforest on Mayotte Island. Brown lemurs have adapted rather well to degraded and secondary forests.

behavior

Brown lemurs live in social groups of between three and 12 individuals, and up to 29 on Mayotte Island. The Mayotte groups are unstable, individuals freely circulating throughout groups every day. Brown lemur groups are unusually peaceful internally, with little or no dominance hierarchy and little dominance of females toward males. Even though abutting territories show a great deal of overlap, groups of neighboring territories make efforts to avoid each other. A brown lemur group holds a territory that can range from 17–50 acres (7–20 ha). Groups often break up into subgroups during the day, re-forming the original large group by nightfall.

feeding ecology and diet

Brown lemur diet consists mostly of fruit, young leaves, and flowers. Diet varies depending on local habitat. Brown lemurs in the southwest make the leaves of the tamarind tree (Tamarindus indica) their staple, while browns on Mayotte eat mostly fruit.

reproductive biology

Polygamous. Reproduction in brown lemurs and development of the young follows the standard Lemuridae pattern. Adults mate April through June and young are born from September through November after a gestation period of about 120 days. A mother usually gives birth to only a single young per season, becoming pregnant annually. Young are weaned in five and a half months and reach sexual maturity at 1.5–2 years of age.

conservation status

Four subspecies, L. f. albocollaris, L. f. collaris, L. f. mayottensis, and L. f. sanfordi are considered Vulnerable. Forest destruction and hunting are the main threats to brown lemurs.

significance to humans

Brown lemurs are hunted and trapped for food.

Crowned lemur

Lemur coronatus

taxonomy

Lemur coronatus Gray, 1842, Madagascar.

other common names

French: Lémur couronné; German: Kronenmaki; Spanish: Lemur coronado.

physical characteristics

Head-and-body length averages 13.4 in (34 cm), tail length 17.7 in (45 cm). Adult body weight averages 4.5 lb (2 kg). The species is sexually dichromatic. Males are gray-brown, females are more gray. The male wears a triangular patch of black fur between his ears, and the female keeps a bright red-orange V-shaped patch leading from the brows back and along the bases of the ears; these are the "crowns" of the common name. Crowned lemurs have scent glands for marking territory on their hands, arms, face, genitals, and anus.

distribution

Crowned lemurs live from near the very northern tip of Madagascar, westward to the Ankarana Massif, and eastward to the east to the Fanambana River south of Vohimarina, in all the coastal zones and at low altitude. Their range includes the slopes of Montagne d'Ambre and the dry forests of the Cap d'Ambre.

habitat

Crowned lemurs have adapted quite well to humid tropical rainforest, dry tropical forests, and forested land in various stages of degradation. The primary humid forest that is home to crowned lemurs is on and around Montagne d'Ambre outside of Antsiranana (Diego Suarez). That forest abuts on the drier forest of the Cap d'Ambre, which is also home to crowned lemurs. The species also lives in remaining areas of lowland dry forest in northern Madagascar. They have adapted well to degraded forest and even grassland studded with isolated trees, the lemurs hiking overland between trees, which they use as refuges.

behavior

Crowned lemur group size runs from two to 11. A typical group has five or six individuals, two adult pairs and one or two younger sub-adults.

Times of activity vary throughout the range. Generally, groups rouse near dawn and forage until nightfall with a four-hour rest in the middle of the day, although some groups will continue to forage for a few hours after night sets in.

feeding ecology and diet

The crowned lemur's menu includes flowers, fruits, and leaves, exact proportions varying with seasons. Fruit is the major food type and may make up almost all of the diet at the end of the dry season, while leaves become important in the rainy season. Crowned lemurs visit water sources during the dry season, even hiking to sources in caves.

reproductive biology

Polygamous. Mating, gestation, birth, and development of the young follow the standard Lemuridae pattern. Births begin in mid-September, earlier for crowned lemur females in humid tropical rainforest, later for females in dryer forests.

conservation status

Vulnerable. Pressure on the species are brought about by deforestation and hunting.

significance to humans

Crowned lemurs are hunted for food and as pests, since they sometimes help themselves to crops.

Red-bellied lemur

Lemur rubriventer

taxonomy

Lemur rubriventer Geoffroy, 1850, Tamatave, Madagascar.

other common names

French: Lémur à ventre rouge, German: Rotbauchlemur, Rotbauchmaki; Spanish: Lemur de vientre rojo.

physical characteristics

Average adult head-and-body length is 15 in (40 cm), with a tail length of 20 in (50 cm) and an adult body weight of 4.5 lb (2 kg). There is some sexual dichromatism. Both sexes have dark brown pelages and black tails, but males have reddish brown undersides (hence the common name) while females differ with cream-white undersides. Males have prominent white teardrop-shaped marks under the eyes, which females do not have. Males keep a scent gland on the tops of their heads.

distribution

Eastern Madagascar.

habitat

Red-bellies live throughout the belt of middle to high-altitude rainforests that runs north-south in eastern Madagascar.

behavior

Red-bellied lemurs live in family groups of up to five individuals, each led by a bonded monogamous pair, though females are still dominant and lead foraging trips. A typical group keeps a territory of 30–37 acres (12–15 ha). Red-bellied lemurs only rarely show territorial behavior. In most cases, groups of neighboring territories, on meeting, scarcely acknowledge each other and almost never fuss.

feeding ecology and diet

Red-bellied lemurs forage for the fruit, flowers, and leaves of at least 30 species of tropical forest plants.

reproductive biology

Monogamous. Individuals reach sexual maturity at two years of age. Pairs mate during May through June and young are born after a 120-day gestation. Mothers bear one young per season. Raising the young in red-bellied lemurs departs from the standard Lemuridae model: the young at first cling to their mother's undersides, switching to riding on her back in two weeks. From then until five weeks, the young will take clinging rides on both parents, sharing time between the two equally. After the fifth week, females start refusing to carry the young, handing them over to the father to carry constantly, until the young are about 100 days old.

conservation status

Vulnerable. The main threat to red-bellied lemurs is the ongoing destruction of Madagascar's eastern rainforests.

significance to humans

Red-bellied lemurs are hunted and trapped for food.

Resources

Books

Mittermeier, R. A., W. R. Konstant, M. E. Nicoll, and O. Langrand. Lemurs of Madagascar: An Action Plan for their Conservation. Gland, Switzerland: IUCN/SSC Primate Specialist Group, 1992.

Plavcan, J., R. F. Kay, W. L. Jungers, and C. P. van Schaik, eds. Reconstructing Behavior in the Primate Fossil Record. New York: Kluwer Academic/Plenum Publishers, 2002.

Rakotosamimanana, B., H. Rasamimanana, J. U. Ganzhorn, and S. M. Goodman, eds. New Directions in Lemur Studies. New York: Kluwer Academic/Plenum Publishers, 1999.

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

Periodicals

Harrington, J. E. "Diurnal Behavior of Lemur mongoz at Ampijoroa, Madagascar." Folia Primatologica 56 (1978): 39–49.

Tattersall, I. "Group Structure and Activity Rhythm in Lemur mongoz (Primates, Lemuriformes) on Anjouan and Moheli Islands, Comoro Archipelago." Anthropological Papers of the American Museum of Natural History 53, 4 (1976): 369–380.

Wilson, J. M., P. D. Stewart, G. S. Ramangason, A. M. Denning, and M. S. Hutchings. "Ecology and Conservation of the Crowned Lemur, Lemur coronatus, at Ankarana, N. Madagascar." Folia Primatologica 52 (1989): 1–26.

Yoder, A. D., M. Cartmill, M. Ruvolo, K. Smith, and R. Vilgalys. "Ancient Single Origin for Malagasy Primates." Proceedings of the National Academy of Sciences 93 (1996): 5122–5126.

Organizations

Duke University Primate Center. 3705 Erwin Road, Durham, NC 27705 United States. Phone: (919) 489-3364. Fax: (919) 490-5394. E-mail: [email protected] Web site: <http://www.duke.edu/web/primate>

Madagascar Fauna Group, San Francisco Zoo. 1 Zoo Road, San Francisco, CA 94132 United States. E-mail: [email protected] Web site: <http://www.selu.com/bio/mfg/text/conservation.html>

Other

2002 IUCN Red List of Threatened Species (International Union for the Conservation of Nature and Natural Resources). <http://www.redlist.org>

Yoder Lab Page. <http://www.basic.nwu.edu/yoder/yoder.html>

Kevin F. Fitzgerald, BS