Overview

Brief Summary

Biology

Brown lemurs live in multimale-multifemale groups without a noticeable hierarchy, generally numbering from 3 to 12 individuals, with 9 to 12 being the norm (2) (5). Breeding is seasonal with mating usually occurring in May and June. The gestation period is approximately 120 days, with infants born between September and October, at the onset of the rainy season. A single offspring is usual, although twins have been reported (2). Brown lemurs reach sexual maturity between one and three years, and the lifespan in the wild is believed to range between 20 and 25 years (2) (5). This species is cathemeral, meaning it is active at varying times throughout the day and night. Fruit, mature leaves, flowers, bark, sap, soil, insects, centipedes and millipedes form the bulk of this lemur's diet (6).
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Description

This medium-sized lemur has a horizontal posture, which is suited to its predominantly quadrupedal mode of movement (2). These lemurs are also capable of leaping considerable distances, their long furry tails assisting them in maintaining their balance (5). The short, dense coat of both sexes is grey-brown on the upperparts, and paler and slightly greyer on the underparts. The face, muzzle and crown are dark-grey to black, with faint pale eyebrow patches and paler grey-brown fur around the ears, cheeks and underneath the chin. The eyes are a rich orange-red (2).
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Distribution

Range Description

This species is found in Madagascar in three populations: in the west, north of the Betsiboka River, where it lives on the high plateau in scattered forest fragments; in eastern Madagascar, to the north of the Mangoro River to the Onive River (generally north-east of Antananarivo as far as the Ambatovaky Special Reserve); and in an isolated population in Ambohitantely Special Reserve, a small reserve of no more than 3,000 ha. They are also on the island of Mayotte in the Comores, where it apparently was introduced by human agency (Mittermeier et al. 2008).

The western populations can be divided into two subpopulations, a southern one extending from the Betsiboka River/Ankarafantsika National Park to the Maverano River, and a northern one ranging from the Andranomalaza River and Manongarivo Reserve to the Mahavavy du Nord River south of Ambilobe. Animals in the northern reaches of this range may also be found throughout the moister forests of the Sambirano region, as well as on the slopes of the Tsaratanana Massif, and are very similar in coloration to the brown lemurs found on the island of Mayotte (Mittermeier et al. 2008). Ranges from sea-level to 400 m.
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Geographic Range

Eulemur fulvus is found on Madagascar and the Comoro Islands. This species includes five subspecies. They are Eulemur fulvus fulvus (common brown lemurs), Eulemur fulvus albifrons (white-fronted lemurs), Eulemur fulvus collaris (collared lemurs), Eulemur fulvus rufus (red-fronted lemurs), and Eulemur fulvus sanfordi (Sanford's lemurs). They all have their own specific ranges within the larger range shared by the entire species.

Red fronted lemurs (E. f. rufus) are found naturally in western and eastern Madagascar. There is also a small introduced population in Southern Madagascar at the Berenty Private Reserve. Common brown lemurs (E. f. fulvis) are found in northwest portions of Madagascar. White-fronted lemurs (E. f. albifrons) are found throughout most of the remaining northeastern rain forest in Madagascar. Collared lemurs (E. f. collaris) are found in southeastern Madagascar, and Sanford's lemurs (E. f. sanfordi) have a very restricted range in northern Madagascar.

At some points, these subspecies exist sympatrically.

Biogeographic Regions: ethiopian (Native )

Other Geographic Terms: island endemic

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Range

This species has a notably disjunct distribution, found in western Madagascar north of the Betsiboka River, on the high plateau in scattered forest fragments, and in eastern Madagascar to the north of the Mangoro River. It has also been introduced to the island of Mayotte in the Comoros (3).
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Physical Description

Morphology

Physical Description

Brown lemurs, like all true lemurs, have binocular vision and long furry tails. They have a scent gland located at their wrist that is used in olfactory communication. This species is sexually dichromatic - its males and females have different fur patterns.

All the members of this species fall within the broad size range of 2 to 4 kg. This is about the size of a housecat. Each subspecies has its own unique markings on its fur.

Red-fronted brown lemurs have an average weight of about 2.7 kg and their average body length is 40 cm. The tail is about 55 cm long. The males are gray to gray-brown and have a reddish crown. The females are reddish-brown. All red-fronted brown lemurs have pale patches over their eyes.

Common brown lemurs, weigh around 2.6 kg. Their body length is 50 cm, as is their tail length. Both males and females are brown to dark-gray with light beards and dark faces.

White-fronted lemurs have a body weight of 2.3 kg, and body length of 40 cm, and a tail length of greater than 50 cm. Generally, this subspecies is dark brown with a lighter underside. Males have a white or cream colored head, ears and beard.

Collared lemurs are around 2.6 kg, with a body length of 50 cm and a tail length of 50 cm as well. Males are brownish-gray with a dark stripe down the back, a dark tail and tail tip, and a lighter underside. Females have a reddish to brown coat and a gray face. Both sexes have a distinct beard that is reddish-brown in females and cream to reddish-brown in males.

Sanford's lemurs weigh around 2.3 kg and have a body length of 40 cm with a tail length of 50+ cm. They can be distinguished from the other subspecies by their fur. Both sexes are dark brown with a lighter underside. Noses, snouts and the area between the eyes are black, and a dark "T" that connects the eyes and nose dominates the head. Males have white-reddish ear tufts and thick beards, providing them the illusion of a 'ragged mane' around their faces.

Range mass: 2 to 4 kg.

Range length: 40 to 50 cm.

Sexual Dimorphism: sexes colored or patterned differently

Average basal metabolic rate: 4.239 W.

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Ecology

Habitat

Habitat and Ecology

Habitat and Ecology
Occurs in the tropical/subtropical dry forest in the west, and tropical moist lowland and montane forest in the east. Groups vary in size from 3-12 (larger on Mayotte) and home ranges on Madagascar vary from approximately seven to 20 ha (Mittermeier et al. 2008).

Systems
  • Terrestrial
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The habitat for E. fulvus varies slightly for each of the included subspecies. Red-fronted lemurs are found in the canopy of deciduous forests in western and eastern Madagascar. Common brown lemurs and collared lemurs live in scattered forest fragments in the high plateaus of western Madagascar. White-fronted lemurs are found in rain forest fragments. Sanford's lemurs inhabit a very limited area of secondary forest.

As mentioned earlier, the habitats for these subspecies do overlap, since some of the groups exist sympatrically.

Terrestrial Biomes: forest ; rainforest

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Found in rainforest, moist montane forest and dry deciduous forest (2). Brown lemurs are arboreal and spend most of their time in the upper layers of the canopy (6).
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Trophic Strategy

Food Habits

This species is largely folivorous. It also eats flowers, fruit, and bark. The diet of E. fulvus varies slightly between subspecies and populations.

Red-fronted lemurs are mainly folivorous (leaf-eating). They also consume pods, stems, flowers, fruit, bark and sap of the kily tree (Tamarindus indica). However, they have very adaptable diets. These lemurs have the ability to shift their normally herbivorous diet to invertebrates and fungi when plant matter is scarce. Eastern populations are specifically known to include insects, bird eggs, and dirt in their diets. They are known to have higher dietary diversity than those populations found in the west and a unique predominance of fruit.

Common brown lemurs, white-fronted lemurs, and collared lemurs eat mainly fruit, young leaves and flowers. Sanford's lemurs feed on primarily fruit, occasional plant parts and invertebrates.

Animal Foods: eggs; insects

Plant Foods: leaves; wood, bark, or stems; fruit; flowers; sap or other plant fluids

Other Foods: fungus

Primary Diet: herbivore (Folivore , Frugivore )

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Associations

Ecosystem Roles

As frugivores, these lemurs are likely to aid in dispersal of seeds. As predators on insects and bird nests, they may affect relevant populations of animals. To the extent that these lemurs serve as prey for other species, they may have some impact on predator populations.

Ecosystem Impact: disperses seeds

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Predation

Predators of these lemurs have not been reported, although possibilities include fossas, raptors, and humans.

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Life History and Behavior

Behavior

Communication and Perception

Grooming is a way for E. fulvus to establish and maintain social bonds. Their unique method of grooming is a result of their 6 lower procumbent teeth that form a dental comb. This instrument is used to groom their own fur and that of the other members in the group.

Communication is achieved by both olfactory and vocal means. Olfactory communication is extremely important and is made possible by the scent glands located at the wrist throat. This type of communication is used for transmitting physical state, location, and individual recognition.

The sounds brown lemurs use for vocal communication have been described and partly deciphered. A nasal sound used in maintaining group cohesion has been described as 'ohn'. A 'cree' or high pitched sound is used as a territorial call, and 'Crou' is the alarm call of this lemur.

In addition to these forms of communication, body postures and facial expressions are likely to be important visual signals.

Communication Channels: visual ; tactile ; acoustic ; chemical

Other Communication Modes: scent marks

Perception Channels: visual ; acoustic

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Life Expectancy

Lifespan/Longevity

One individual of this species lived over 36 years in captivity. It is likely that, as with other lemurs in the genus, the maximum lifespan in the wild ranges between 20 and 25 years.

Range lifespan

Status: captivity:
36 (high) years.

Typical lifespan

Status: wild:
25 (high) years.

Average lifespan

Status: captivity:
35.5 years.

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Lifespan, longevity, and ageing

Maximum longevity: 35.5 years (captivity) Observations: One specimen lived 35.5 years in captivity. Assuming it was received as an adult, it could have lived to the age of 37 or even more because its exact date of death is unknown. A hybrid between a brown and a black lemur reportedly lived 39.3 years (Richard Weigl 2005).
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Reproduction

The mating system of these lemurs has not been reported. However, other species in the genus Eulemur are either monogamous or polygynous. It is likely that E. fulvus is similar. Unlike other members of the genus, females are not usually dominant to males, so the degree to which females exert active mate choice is not known.

Brown lemurs reach sexual maturity between 1 and 2 years of age. Their mating habits are very seasonal with mating occuring sometimes in late May (Sanford's lemurs only) but usually throughout June and July (all subspecies). The gestation period for these animals is approximately 120 days. Infants are born in the fall, between September and November. Only one infant is born per year to each mother.

Breeding interval: These lemurs breed once per year.

Breeding season: Mating occurs sometimes in late May (Sanford's lemurs only) but usually throughout June and July (all subspecies)

Average number of offspring: 1.

Average gestation period: 120 days.

Range weaning age: 4 to 6 months.

Range age at sexual or reproductive maturity (female): 1 to 2 years.

Range age at sexual or reproductive maturity (male): 1 to 2 years.

Key Reproductive Features: iteroparous ; seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); viviparous

Average birth mass: 83.3 g.

Average gestation period: 118 days.

Average number of offspring: 1.1.

For the first three weeks of their lives, young lemurs hang onto the mother's bellies. They alter their grasp only to nurse. After three weeks have passed, they shift and ride on the mother's backs. They then begin to take their first steps. Following this, they start to sample solid food, nibbling on whatever the other members of the group happen to be eating. This is their first sign of independence. Nursing continues but its importance in the infant's diet tapers. The young lemur is weaned after approximately 4 to 6 months - usually by January.

The role of males in parental care in this species has not been described.

Parental Investment: pre-fertilization (Provisioning, Protecting: Female); pre-hatching/birth (Provisioning: Female, Protecting: Female); pre-weaning/fledging (Provisioning: Female, Protecting: Female); pre-independence (Provisioning: Female, Protecting: Female); extended period of juvenile learning

  • Nowak, R. 1999. Walker's Mammals of the World, Sixth Edition. Baltimore and London: The Johns Hopkins University Press.
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Molecular Biology and Genetics

Molecular Biology

Barcode data: Eulemur fulvus

The following is a representative barcode sequence, the centroid of all available sequences for this species.


There are 4 barcode sequences available from BOLD and GenBank.

Below is a sequence of the barcode region Cytochrome oxidase subunit 1 (COI or COX1) from a member of the species.

See the BOLD taxonomy browser for more complete information about this specimen and other sequences.

AATCGTTGATTCTACTCAACCAATCATAAAGATATCGGAACTCTTTACCTCCTATTTGGGGCTTGAGCAGGCATGGTAGGAACAGCTCTT---AGCCTTTTAATTCGAGCAGAACTTGGTCAACCCGGGGCTTTATTAGGAGAT---GACCAAATCTACAATGTAATCGTAACAGCTCATGCTTTCGTTATAATCTTTTTCATAGTTATACCTATCATAATTGGGGGCTTTGGGAACTGACTAGTTCCCTTAATA---ATTGGAGCCCCTGATATAGCATTCCCTCGAATAAATAATATAAGCTTTTGGCTTCTACCACCATCCTTTCTACTACTTCTAGCATCCTCAATAGTAGAAGCAGGTGCTGGAACTGGGTGGACCGTGTATCCTCCTCTAGCTGGTAATTTAGCCCATGCAGGAGCCTCCGTAGATTTA---ACAATTTTTTCATTACACCTAGCAGGAGTATCCTCAATTCTAGGGGCTATCAACTTTATCACCACAGTAATTAACATAAAACCCCCAGCCATATCACAATATCAAACTCCTCTATTTGTATGATCCGTGATAATTACCGCTGTCCTTTTACTTTTATCCCTACCAGTCCTAGCAGCA---GGAATCACTATACTCTTAACTGACCGCAATCTCAACACTACATTTTTTGATCCCGCAGGAGGAGGTGATCCAATTCTATATCAACACCTATTCTGATTCTTCGGACATCCTGAAGTCTACATCTTAATCCTCCCAGGTTTTGGCATAATTTCTCACATTGTCACATATTATTCAGGTAAAAAA---GAGCCATTTGGTTATATAGGCATAGTCTGAGCTATAATATCTATTGGCTTCTTAGGATTTATCGTATGAGCGCACCATATATTCACAGTAGGTATAGACGTAGATACTCGAGCATACTTTACGTCTGCCACTATAATTATTGCTATTCCCACTGGTGTAAAAGTATTTAGCTGACTA---GCAACACTACATGGCGGA---AACATCAAATGGTCACCCGCTATACTATGAGCCCTTGGCTTTATTTTCCTATTCACGGTTGGAGGCTTAACAGGAATTGTACTTGCCAATTCATCACTGGATATTGTTCTCCATGATACTTATTATGTAGTAGCCCACTTCCATTATGTA---CTATCAATAGGAGCAGTCTTTGCTATCATAGGAGGCTTTGTTCACTGATTTCCTTTATTTTCAGGATATACCTTAGACGACACTTGAGCTAAAATTCACTTCTCAATTATATTTGTGGGTGTAAACATAACCTTTTTCCCACAACACTTCTTAGGTTTATCTGGAATACCCCGA---CGCTATTCCGATTACCCGGACGCCTACACC---ATATGAAATACTGTTTCATCCATCGGCTCTTTCATCTCCCTAACAGCAGTTATATTAATAGTTTTCATGATTTGGGAGGCCTTTGCCTCAAAACGAGAGGTC---CTAATAGTGGAACTCACACCAACTAAC
-- end --

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Statistics of barcoding coverage: Eulemur fulvus

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 4
Specimens with Barcodes: 6
Species With Barcodes: 1
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Conservation

Conservation Status

IUCN Red List Assessment


Red List Category
NT
Near Threatened

Red List Criteria

Version
3.1

Year Assessed
2008

Assessor/s
Andrainarivo, C., Andriaholinirina, V.N., Feistner, A., Felix, T., Ganzhorn, J., Garbutt, N., Golden, C., Konstant, B., Louis Jr., E., Meyers, D., Mittermeier, R.A., Perieras, A., Princee, F., Rabarivola, J.C., Rakotosamimanana, B., Rasamimanana, H., Ratsimbazafy, J., Raveloarinoro, G., Razafimanantsoa, A., Rumpler, Y., Schwitzer, C., Thalmann, U., Wilmé, L. & Wright, P.

Reviewer/s
Mittermeier, R.A. & Rylands, A.B. (Primate Red List Authority)

Contributor/s

Justification
Listed as Near Threatened as the species is thought to have undergone a reduction of 20-25% over the past 24 years (assuming a generation length of 8 years) due primarily to a decline in area and quality of habitat within the known range of the species and due to known levels of exploitation. Almost qualifies as threatened under criterion A2cd.

History
  • 1994
    Vulnerable
    (Groombridge 1994)
  • 1994
    Rare
    (Groombridge 1994)
  • 1990
    Rare
    (IUCN 1990)
  • 1990
    Rare
    (IUCN 1990)
  • 1990
    Vulnerable
    (IUCN 1990)
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The conservation status of brown lemurs is partially due to their restricted geographical area. Their primary threat is habitat destruction. Habitat destruction is largely the result of the explosive growth rate of the human population of Madagascar. This species is placed in a somewhat lower risk category (Vulnerable) because of its presumably large wild population and occurrence in a number of protected areas.

For red-fronted lemurs, the western habitats are largely at risk because of burning and clearing of land for pasture. In the east, the chief hazards are the slash-and-burn agriculture and forest cutting for fuel wood and construction. This subspecies occurs in some protected areas of Madagascar and can be found in captivity in 22 zoos worldwide where approximately 100 individuals are held.

Forest destruction is the primary threat to the survival of common brown lemurs. They are also hunted throughout much of their range. This subspecies is found in protected areas in Madagascar, and may be one of the lowest risk subspecies of brown lemur. Common brown lemurs have bred in captivity and there are currently about 140 animals at 40 institutions worldwide.

The white-fronted lemurs are threatened by the destruction of Madagascar's eastern rain forest for slash-and-burn agriculture. This animal is also hunted for food throughout most of its range. Presently, it does exist in protected areas in Madagascar. White-fronted lemurs do breed in captivity, and there are over 200 animals in captivity at more than 40 zoos worldwide.

Forest destruction is the primary threat to the survival of collared lemurs as well. They too, are hunted for food throughout much of their range. In addition to this, they are occasionally trapped for the pet trade. Collared lemurs occur naturally in only one of Madagascar's protected areas, but have been introduced into two others. Approximately 40 collared lemurs are currently in captivity in 6 institutions worldwide.

Sanford's lemurs are also threatened by forest destruction. This subspecies, however, does appear to be able to survive in slightly degraded habitats. Although Sanford's lemurs are found in protected areas in Madagascar, the level of protection varies among the reserves. This is because poaching and brush fires are common events in many of Madagascar's nature reserves. There are only two captive breeding groups of sanford's lemurs.

US Federal List: endangered

CITES: appendix i

IUCN Red List of Threatened Species: near threatened

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Status

Recent scientific thought is that the six subspecies of brown lemur previously acknowledged should be elevated to individual species status, with the nominate subspecies, the common brown lemur (E. f. fulvus), being categorised as the full species, the brown lemur (Eulemur fulvus) (3). As a subspecies, the common brown lemur (E. f. fulvus) is classified as Lower Risk/near threatened (LR/nt) on the IUCN Red List 2004 (1). Listed under Appendix I of CITES (4).
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Population

Population
Population densities range from 40-60 individuals/km² (Mittermeier et al. 2008).

Population Trend
Decreasing
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Threats

Major Threats
Forest destruction, due primarily to slash-and-burn practices, charcoal production and illegal logging, is the principal threat, but hunting is increasingly becoming a significant threat (including with blowpipes, firearms, bow-and-arrows and traps) and sometimes entire groups are captured.
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Habitat destruction remains the primary threat to the brown lemur, largely as a result of the explosive growth in the human population on Madagascar (5). Eastern areas of rainforest are destroyed by slash-and-burn agriculture and by forest cutting for fuel wood and construction, while dryer western forests are cleared by fires started to promote new flushes of pasture for grazing cattle. Hunting and trapping for food or the pet trade may also constitute a threat to the brown lemur in some parts of its range (2) (5).
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Management

Conservation Actions

Conservation Actions
This species is listed on Appendix I of CITES. This species is reported to occur in four national parks (Ankarafantsika, Mantadia, Andringitra, and Zahamena), two strict nature reserves (Tsaratanana and Zahamena), and seven special reserves (Ambatovaky, Ambohitantely, Analamazaotra, Bora, Mangerivola, Manongarivo, and Tampoketsa-Analamaitso) (Mittermeier et al. 2008)). There is a relatively large worldwide captive population.
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Conservation

The brown lemur is found in at least 13 protected areas, including four national parks, two strict nature reserves and seven special reserves (3). Captive bred populations also exist in institutions worldwide (5). The fate of the brown lemur in the wild will most probably be determined by the future of its forest habitat, which needs to be better preserved if the survival of this lemur is to be safeguarded.
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Relevance to Humans and Ecosystems

Benefits

Economic Importance for Humans: Negative

Eulemur fulvus has no known negative effects on humans.

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Economic Importance for Humans: Positive

The protected areas of Madagascar, which many lemurs (as well as many other types of flora and fauna) reside in, have become quite an attraction for tourists. Communities in Madagascar benefit greatly from this. They receive fifty percent of national park entry fees. Local inhabitants also benefit by serving as guides and by selling handicrafts to the tourists.

Members of this species are sometimes hunted for meat.

Positive Impacts: food ; ecotourism

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Wikipedia

Common brown lemur

The common brown lemur (Eulemur fulvus), or brown lemur, is a species of lemur in the family Lemuridae. It is found in Madagascar and Mayotte.[2]

Range[edit]

The common brown lemur lives in western Madagascar north of the Betsiboka River and eastern Madagascar between the Mangoro River and Tsaratanana, as well as in inland Madagascar connecting the eastern and western ranges.[3] They also live on the island of Mayotte, although this population is believed to have been introduced there by man.[3]

Physical description[edit]

The common brown lemur has a total length of 84 to 101 centimeters, including 41 to 51 centimeters of tail.[4] Weight ranges from 2 to 3 kilograms.[4] The short, dense fur is primarily brown or grey-brown.[4] The face, muzzle and crown are dark grey or black with paler eyebrow patches, and the eyes are orange-red.[4]

Similar lemur species within their range include the mongoose lemur, E. mongoz, in the west and the red-bellied lemur, E. rubriventer, in the east.[4] They can be distinguished from these species by the fact that E. mongoz is more of a grey color and E. rubriventer is more reddish. There is also some overlap with the black lemur in northeast Madagascar in the Galoko, Manongarivo and Tsaratanana Massifs.[5] There is also overlap and hybridization with the white-fronted brown lemur, E. albifrons, in the northeast portion of the common brown lemur's range.[6]

Diet[edit]

The common brown lemur's diet consists primarily of fruits, young leaves, and flowers.[3] In some locations it eats invertebrates, such as cicadas,[4] spiders[4] and millipedes.[7] It also eats bark, sap, soil and red clay (see geophagy).[7] It can tolerate greater levels of toxic compounds from plants than other prosimians can.[3][7]

Behavior[edit]

Consistent with its large range, the common brown lemur occupies a variety of forest types, including lowland rainforests, montane rainforests, moist evergreen forests and dry deciduous forests.[4] They spend about 95% of their time in upper layers of the forest and less than 2% of their time on the ground.[7]

They normally live in groups of 5 to 12, but group size can be larger, especially on Mayotte.[4] Groups occupy home ranges of 1 to 9 hectares in the west, but more than 20 hectares in the east.[8] Groups include members of both sexes, including juveniles, and there are no discernible dominance hierarchies.[4]

They are primarily active during the day, but can exhibit cathemeral activity and continue into the night, especially during full moons[4] and during the dry season.[3][9]

In the western part of its range, the common brown lemur overlaps that of the mongoose lemur, and the two species sometimes travel together.[7] In the areas of overlap, the two species also adapt their activity patterns to avoid conflict.[9] For example, the Mongoose Lemur can become primarily nocturnal during the dry season in the areas of overlap.

At Berenty (south Madagascar) there is a population of introduced E. fulvus rufus x collaris.[10] These lemurs show linear hierarchy, adult female dominance, and the presence of conciliatory behavior after aggressions.[11] Additionally, stress levels (measured via self-directed behaviors) decrease at the increase of the hierarchical position of individuals within the social group and reconciliation is able to bring stress down to the baseline levels.[12]

Reproduction[edit]

The common brown lemur's mating season is May and June.[4] After a gestation period of about 120 days, the young are born in September and October.[4] Single births are most common, but twins have been reported.[4] The young are weaned after about 4 to 5 months.[4][7] Sexual maturity occurs at about 18 months,[4] and females give birth to their first young at 2 years old.[7] Life span can be as long as 30+ years.[7]

Taxonomy[edit]

Five additional currently recognized species of lemur were until 2001 considered subspecies of E. fulvus.[13] These are:

However, a number of zoologists believe that E. albifrons and E. rufus should continue to be considered subspecies of E. fulvus.[13]

References[edit]

  1. ^ Groves, C. P. (2005). Wilson, D. E.; Reeder, D. M, eds. Mammal Species of the World (3rd ed.). Baltimore: Johns Hopkins University Press. p. 115. OCLC 62265494. ISBN 0-801-88221-4. 
  2. ^ a b Andriaholinirina, N., Baden, A., Blanco, M., Chikhi, L., Cooke, A., Davies, N., Dolch, R., Donati, G., Ganzhorn, J., Golden, C., Groeneveld, L.F., Hapke, A., Irwin, M., Johnson, S., Kappeler, P., King, T., Lewis, R., Louis, E.E., Markolf, M., Mass, V., Mittermeier, R.A., Nichols, R., Patel, E., Rabarivola, C.J., Raharivololona, B., Rajaobelina, S., Rakotoarisoa, G., Rakotomanga, B., Rakotonanahary, J., Rakotondrainibe, H., Rakotondratsimba, G., Rakotondratsimba, M., Rakotonirina, L., Ralainasolo, F.B., Ralison, J., Ramahaleo, T., Ranaivoarisoa, J.F., Randrianahaleo, S.I., Randrianambinina, B., Randrianarimanana, L., Randrianasolo, H., Randriatahina, G., Rasamimananana, H., Rasolofoharivelo, T., Rasoloharijaona, S., Ratelolahy, F., Ratsimbazafy, J., Ratsimbazafy, N., Razafindraibe, H., Razafindramanana, J., Rowe, N., Salmona, J., Seiler, M., Volampeno, S., Wright, P., Youssouf, J., Zaonarivelo, J. & Zaramody, A. (2014). "Eulemur fulvus". IUCN Red List of Threatened Species. Version 2014.1. International Union for Conservation of Nature. Retrieved 2014-06-16.  Listed as Near Threatened (NT v3.1)
  3. ^ a b c d e Russell Mittermeier et al. (2006). Lemurs of Madagascar (2nd ed.). pp. 272–274. ISBN 1-881173-88-7. 
  4. ^ a b c d e f g h i j k l m n o p Nick Garbutt (2007). Mammals of Madagascar. pp. 155–156. ISBN 978-0-300-12550-4. 
  5. ^ Russell Mittermeier et al. (2006). Lemurs of Madagascar (2nd ed.). p. 288. ISBN 1-881173-88-7. 
  6. ^ Russell Mittermeier et al. (2006). Lemurs of Madagascar (2nd ed.). p. 282. ISBN 1-881173-88-7. 
  7. ^ a b c d e f g h Noel Rowe (1996). The Pictorial Guide to the Living Primates. p. 40. ISBN 0-9648825-0-7. 
  8. ^ name=perspective>Lisa Gould and Michelle Sauther (2007). "Lemuriformes". In Christina J. Campbell, Agustin Fuentes, Katherine C. MacKinnon, Melissa Panger and Simon K. Bearder. Primates in Perspective. p. 53. ISBN 978-0-19-517133-4. 
  9. ^ a b Robert W. Sussman (1999). Primate Ecology and Social Structure Volume 1: Lorises, Lemurs and Tarsiers. pp. 186–187. ISBN 0-536-02256-9. 
  10. ^ Alison Jolly, Naoki Koyama, Hantanirina Rasamimanana, Helen Crowley, and George Williams (2006). "Berenty Reserve: a research site in southern Madagascar". In A. Jolly, R. W. Sussman, N. Koyama & H. Rasamimanana. Ringtailed Lemur Biology: Lemur catta in Madagascar. pp. 32–42. ISBN 0-387-32669-3. 
  11. ^ Norscia, I., Palagi, E. (2010). "Do wild brown lemurs reconcile? Not always". Journal of Ethology. doi:10.1007/s10164-010-0228-y. 
  12. ^ Palagi, E., Norscia, I. (2010). "Scratching around stress: hierarchy and reconciliation make the difference in wild brown lemurs Eulemur fulvus". Stress. doi:10.3109/10253890.2010.505272. 
  13. ^ a b Russell Mittermeier et al. (2006). Lemurs of Madagascar (2nd ed.). p. 251. ISBN 1-881173-88-7. 
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