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Lifespan, longevity, and ageing

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Maximum longevity: 36.2 years (captivity) Observations: A hybrid between a brown and a black lemur lived 39 years (Ronald Nowak 1999). One captive female was still alive after 36.2 years in captivity (Richard Weigl 2005).
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Life Expectancy

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These animals are reported to live 20 to 25 years, presumably in captivity.

Range lifespan
Status: captivity:
20 to 25 years.

Average lifespan
Status: captivity:
39.0 years.

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Lundrigan, B. and D. Davis 2000. "Eulemur macaco" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Eulemur_macaco.html
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Barbara Lundrigan, Michigan State University
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Daniel Davis, Michigan State University
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Nancy Shefferly, Animal Diversity Web
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Habitat

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Black lemurs are tree dwellers. They can be found in four habitats: primary rain forest, secondary forest, timber plantations and food crop plantations.

Habitat Regions: tropical ; terrestrial

Terrestrial Biomes: rainforest

Other Habitat Features: agricultural

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Lundrigan, B. and D. Davis 2000. "Eulemur macaco" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Eulemur_macaco.html
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Barbara Lundrigan, Michigan State University
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Daniel Davis, Michigan State University
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Nancy Shefferly, Animal Diversity Web
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Distribution

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Black lemurs are limited to the northwestern tip of Madagascar and the two adjacent islands of Nosy Komba and Nosy Be. In Madagascar, the two subspecies are separated by the Andranomalaza river, but clear separation only occurs in a relatively small area (Rabarivola and Meyers,1991). Eulemur macaco macaco can be found to the north of the river and Eulemur macaco flavifrons to the south of it (Rabarivola and Meyers, 1991).

Biogeographic Regions: ethiopian (Native )

Other Geographic Terms: island endemic

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Lundrigan, B. and D. Davis 2000. "Eulemur macaco" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Eulemur_macaco.html
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Barbara Lundrigan, Michigan State University
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Daniel Davis, Michigan State University
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Nancy Shefferly, Animal Diversity Web
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Trophic Strategy

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During the rainy season, their diet seems to consist mainly of fruit. They have also been reported to eat mushrooms and millipedes on occasion during this season. Early in the dry season, a significant part of their diet comes from the nectar of flowers. Other things included in the dry season diet are seed pods, leaves, and flowers (Kappeler and Ganzhorn, 1993).

Animal Foods: terrestrial non-insect arthropods

Plant Foods: leaves; seeds, grains, and nuts; fruit; nectar; flowers

Other Foods: fungus

Primary Diet: herbivore (Folivore , Nectarivore )

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Lundrigan, B. and D. Davis 2000. "Eulemur macaco" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Eulemur_macaco.html
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Barbara Lundrigan, Michigan State University
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Daniel Davis, Michigan State University
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Nancy Shefferly, Animal Diversity Web
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Associations

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As frugivores, these lemurs are likely to play some role in seed dispersal. However, because they eat nectar, they may also be important pollinators. To the extent that these lemurs are food for predators, they may have impact in local food webs.

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Lundrigan, B. and D. Davis 2000. "Eulemur macaco" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Eulemur_macaco.html
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Barbara Lundrigan, Michigan State University
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Daniel Davis, Michigan State University
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Nancy Shefferly, Animal Diversity Web
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Benefits

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Humans have found the black lemurs to be useful as a food resource and for their furs. They can also be trapped and sold to people as pets or used as attractions in zoos.

Positive Impacts: pet trade ; food ; body parts are source of valuable material; research and education

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Lundrigan, B. and D. Davis 2000. "Eulemur macaco" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Eulemur_macaco.html
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Barbara Lundrigan, Michigan State University
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Daniel Davis, Michigan State University
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Nancy Shefferly, Animal Diversity Web
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Benefits

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Black lemurs are considered crop pests by farmers in some areas.

Negative Impacts: crop pest

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Lundrigan, B. and D. Davis 2000. "Eulemur macaco" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Eulemur_macaco.html
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Barbara Lundrigan, Michigan State University
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Daniel Davis, Michigan State University
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Nancy Shefferly, Animal Diversity Web
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Conservation Status

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All lemurs are considered endangered by the U.S. Fish and Wildlife Service and are protected under Appendix I of CITES. They are also listed as vulnerable in the IUCN Red Data Book. Evidence indicates that their numbers are declining. The main threats to black lemurs include habitat destruction, poaching for their meat or fur, and capture for the pet trade or zoos. They have also been killed in some areas as crop pests.

Lemurs breed fairly well in captivity and are popular in zoos worldwide. The St. Louis Zoological Park in the United States coordinates the Species Survival Plan (SSP) for black lemurs. Reintroduction of lemurs bred in captivity may be used one day to boost wild populations.

US Federal List: endangered

CITES: appendix i

IUCN Red List of Threatened Species: vulnerable

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Lundrigan, B. and D. Davis 2000. "Eulemur macaco" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Eulemur_macaco.html
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Barbara Lundrigan, Michigan State University
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Daniel Davis, Michigan State University
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Nancy Shefferly, Animal Diversity Web
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Behavior

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These animals use vocalizations, scent marks, body postures and facial expressions, as well as tactile communication.

Communication Channels: visual ; tactile ; acoustic ; chemical

Other Communication Modes: scent marks

Perception Channels: visual ; tactile ; acoustic ; chemical

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Lundrigan, B. and D. Davis 2000. "Eulemur macaco" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Eulemur_macaco.html
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Barbara Lundrigan, Michigan State University
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Daniel Davis, Michigan State University
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Nancy Shefferly, Animal Diversity Web
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Untitled

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Black lemurs may breed with blue-eyed lemurs and will produce brown-eyed offspring in every case (Duke University, 1998).

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Lundrigan, B. and D. Davis 2000. "Eulemur macaco" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Eulemur_macaco.html
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Barbara Lundrigan, Michigan State University
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Daniel Davis, Michigan State University
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Nancy Shefferly, Animal Diversity Web
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Morphology

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Black lemurs are primitive primates that are about the size of a house cat. Adults can weigh about 2.4 kg (Duke Primate Center, 1998). Head and body lengths vary between 300 and 450 mm (Nowak, 1999). There are two subspecies of the black lemur: black lemurs (Eulemur macaco macaco) and blue-eyed lemurs (Eulemur macaco flavifrons). These subspecies are similar in size, shape and behavior. Differences between the two subspecies include habitat, coat and eye color.

There is striking sexual dimorphism in color. Males in both subspecies are black. Female black lemurs have a dark coat which lightens to a deep rust on the sides. They are off-white on the stomachs. Female blue-eyed lemurs have a coat that is reddish-tan in color over the entire body. All black lemurs have brown eyes as opposed to blue-eyed lemurs which all have turquoise blue eyes (Duke University, 1998).

Range mass: 2 to 2.5 kg.

Average mass: 2.5 kg.

Range length: 300 to 450 mm.

Sexual Dimorphism: sexes colored or patterned differently; female more colorful

Other Physical Features: endothermic ; bilateral symmetry

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Lundrigan, B. and D. Davis 2000. "Eulemur macaco" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Eulemur_macaco.html
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Barbara Lundrigan, Michigan State University
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Daniel Davis, Michigan State University
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Nancy Shefferly, Animal Diversity Web
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Associations

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Predators of these lemurs are not known. However, it seems likely that animals like fossas and raptors are possible predators.

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Lundrigan, B. and D. Davis 2000. "Eulemur macaco" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Eulemur_macaco.html
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Barbara Lundrigan, Michigan State University
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Daniel Davis, Michigan State University
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Nancy Shefferly, Animal Diversity Web
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Reproduction

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The mating system of these animals has not been well studied. In the wild, groups range in size from 4 to 15 individuals. Females are dominant to males, and there may be some exchange of adults between groups. In captivity, females have their choice of mates. From this information, it seems likely that breeding is polygynous.

Mating System: polygynous

Black lemurs breed seasonally in June and July. Birth occurs after a gestation period of 120 to 129 days. One offspring is usually born, however, twins are fairly common. The young are weaned at five to six months of age. Sexual maturity is reached by about 2 years of age.

Breeding interval: These animals can breed annually.

Breeding season: Breeding occurs in June and July.

Range number of offspring: 1 to 2.

Range gestation period: 120 to 129 days.

Range weaning age: 5 to 6 months.

Average age at sexual or reproductive maturity (female): 2 years.

Average age at sexual or reproductive maturity (male): 2 years.

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

Average birth mass: 74.67 g.

Average number of offspring: 1.

Average age at sexual or reproductive maturity (male)
Sex: male:
548 days.

Average age at sexual or reproductive maturity (female)
Sex: female:
548 days.

Infants can be found clinging to their mothers' bellies for the first 3 weeks and will shift only to nurse. At about 3 weeks of age, the young lemur will begin to ride on its mother's back and will soon after take its first tentative steps. Nursing continues until about 5 to 6 months of age. Mothers provide grooming, protection, and transportation for their young, as well as valuable socialization. The role of males in parental care is not clear, although there is some evidence that males can be infanticidal in captivity. They may, therefore, have some role in protecting their offspring in the wild.

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

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Lundrigan, B. and D. Davis 2000. "Eulemur macaco" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Eulemur_macaco.html
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Barbara Lundrigan, Michigan State University
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Daniel Davis, Michigan State University
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Nancy Shefferly, Animal Diversity Web
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Biology

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Black lemurs live in groups of between 2-15 individuals, with adult males and females in equal numbers, together with their dependant offspring (5). The activities and movements of the group are dictated by the dominant female, and group relationships are maintained by grunts, contact calls and grooming. Home ranges extend for five to six hectares and there is considerable overlap with the ranges of neighbouring groups (2). This species, like other lemurs, exhibits an activity pattern that is virtually unique amongst primates and rarely encountered amongst other arboreal mammals. They have activity bursts which may occur during the day and the night, though most activity occurs in the early morning and late afternoon (5). This activity pattern is called cathemeral, meaning 'all hours', contrasting with the usual distinction between nocturnal and diurnal (9). Foraging is concentrated in the middle and upper parts of the canopy, where this lemur feeds on fruit, flowers, leaves, fungi and occasionally invertebrates like millipedes. This primate plays an extremely important role is seed dispersal through the forests because it has such a high amount of fruit in its diet (6). In the dry season nectar becomes an important part of its diet as well (2). During the day it forages in the understory of the canopy where it is more protected from predatory birds such as hawks, and at night is able to feed in the upper levels (2). The breeding season is seasonal and births occur after a gestation period of 12 - 129 days. One offspring is usual, though twins are fairly common. The young cling to their mother's belly for three weeks and will only move to suckle. After three weeks the young are heavier and ride on the mother's back, and at 5 - 6 months of age they are fully weaned (5).
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Conservation

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The back lemur occurs in three protected areas in Madagascar but its range is limited and its distribution patchy (2). Sclater's lemurpopulations are even lower as this subspecies seems less adaptable to the disturbed forest (2). A national park has been proposed within this lemur's range, and it is hoped that this will allow this species to recover (2). However, the threats facing Madagascar's forests are so great it seems unlikely the black lemur will recover without intervention. Captive breeding projects in association with the Species Survival Plan are now being co-ordinated by the St. Louis Zoological Park in the United States. Fortunately lemurs breed well in captivity and it is hoped that reintroducing these lemurs may be successful and boost wild populations in the near future (7).
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Description

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There are two subspecies of the black lemur, the black lemur Eulemur macaco macaco and the Sclater's or blue eyed lemur E. m. flavifrons. These subspecies are similar in size, shape and behaviour, though they differ in their coat and eye colour and their habitat preferences (2). In both subspecies the sexes look quite different from each other. Male black lemurs have a dark brown to black pelage with black tufted ears and beady yellow-orange eyes, while females look so different they were thought for a long time to be a different species (4). Black lemur females are tawny on the back and head; their underparts are golden-brown to rich-chestnut brown, with paler fur on the limbs, and the tail is a darker chestnut brown (2). The females' ears are also dramatically tufted, but with long white hair which extends around the cheeks (3). Male Sclater's lemursnormally have a completely black pelage, which is shorter in length and has a softer appearance than that of the black lemur. The forehead has a distinct ridge of hair that forms a distinctive crest, and the ears are not tufted (2). As the alternative name of the Sclater's lemur, the blue-eyed black lemur, suggests, this subspecies has blue eyes, which is highly unusual in primates and very striking (2). Females of this subspecies have paler brown coloured underparts and dark grey hands and feet. Their crown is reddish-tan in colour and they have short white beards which may have reddish tinges (2).
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Habitat

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The habitats of these two subspecies differ. The black lemuroccurs in moist Sambirano forests, rainforests on offshore islands, and in modified habitats of timber, coffee and cashew nut plantations, while Sclater's lemur occurs in mixed moist forests, dry deciduous forests and plantations (2).
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Range

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Like all lemurs, these two subspecies are endemic to Madagascar, a large island off the coast off east Africa. They occur in the forests of the Sambirano region on the north-western tip of the island as well as on the islands of Nosy Be and Nosy Komba. These subspecies are separated by the Andranomalaza River, though clear separation occurs only in a small area. The black lemur occurs to the north of the river, and Sclater's lemur to the south (4).
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Status

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The black lemur is classified as Vulnerable (VU A1cd) on the IUCN Red List (1) and is listed on Appendix I on CITES (8). Two subspecies of the black lemur are recognised. The black lemur Eulemur macaco macaco is classified as Vulnerable (VU B1+2bc), and Sclater's lemur (or blue-eyed black lemur) E. m. flavifrons is classified as Critically Endangered (CR A1cd, B1+2bc) on the IUCN Red List (1) (3).
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Threats

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This species is threatened by habitat destruction, hunting for meat or fur, and by trapping for the pet trade (2).
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Habitat

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E. flavifrons lives in fragmented areas of primary and secondary forest. Primary forest contains more densely packed trees and less human disturbance, whereas secondary forest is less dense and has had more human disturbance, such as logging (Schwitzer et al., 2007b). The primary forest fragments have many benefits for E. flavifrons. The many trees that have been left undisturbed in the primary forests are often very tall, about 10-15 m. This allows E. flavifrons to spend a large portion of their day at high elevations in the trees, especially when they are sleeping and eating (Schwitzer et al., 2007b). Likewise, tall, old trees are often unaffected by storms and high winds on the island (Volampeno et al., 2013). Also, a higher diversity of trees in primary forests allows for more food diversity for feeding and more choices for sleeping trees (Schwitzer et al., 2007b). When species move within the primary forests, there are many options to choose from for food and shelter because there is so much variety (Volampeno et al., 2013).

Tree density also plays a large role in E. flavifrons habitat preferences (Seiler et al., 2013). When tree density is higher, there tend to be more individuals. This correlation may arise because trees provide refuge from predators as well as increased food availability (Seiler et al., 2013). A study by Schwitzer et al. (2012) showed that the secondary forest fragments have fewer individuals, supporting the idea that greater tree density in primary forests plays a large role in habitat selection by E. flavifrons (Schwitzer et al., 2010 and Schwitzer et al., 2012). As E. flavifrons prefers primary forests over secondary forests, they are vulnerable to the destruction of the primary forests (Schwitzer et al., 2007b, Seiler et al., 2013, & Volampeno et al., 2013).

E. flavifrons individuals tend to travel to areas where there is more food, which can be difficult in Madagascar because there are both dry and wet seasons (Randriatahina and Roeder, 2013). The wet season, which is the summer season, falls from December to March. The rainfall during this time is very unpredictable, and includes serious storms and monsoon winds. The rain contributes to a plentiful food supply during the wet season (Volampeno et al., 2011a). In contrast, the dry season contributes to a food shortage from the months of May to September, the winter season, which can be very windy (Volampeno et al., 2011).

E. flavifrons has the shortest distance between lemurs, meaning that they live in a small range of habitat compared to other species. One study of lemur populations surrounding Radama National Park, a dry deciduous foreset, found 29 different groups (about 8 individuals per group), and 228 individuals total, with 97.3 lemurs per Km2. The population was shown to be greater on the inside of the park compared to the outside of the park (Volampeno et al., 2010). This indicates that the lemurs tend to reside inside the national park, where they are safer from human interference.

The two subspecies of Eulemur macaco, E. flavifrons and E. macaco, may share a common habitat hybrid zone, as determined by an ecological niche model, based on climate and type of forest. Both of the species could potentially survive in this habitat hybrid zone. However, the actual habitat locations of the populations for both species do not overlap with each other at this time (Blair et al., 2013).

References

  • Schwitzer, C., G. G. Randriatahina, and S. Volampeno. 2012. Sclater’s black lemur or blue-eyed black lemur, Eulemur flavifrons (Gray 1867). Madagascar.
  • Schwitzer, N., G. H. Randriatahina, W. Kaumanns, D. Hoffmeister, and C. Schwitzer. 2007b. Habitat utilization of blue-eyed black lemurs, Eulemur macaco flavifrons (Gray, 1867), in primary and altered forest fragments. Primate Conserv. 22(1): 79–87.
  • Volampeno, M. S. N., G. Randriatahina, and C. T. Downs. 2013. Structure and Composition of Ankarafa Forest, Sahamalaza-Iles Radama National Park, Madagascar: implications for the frugivorous endemic blue-eyed black lemur (Eulemur flavifrons). S. Afr. J. Wildl. Res. 43(2): 91–102.
  • Volampeno, M. S. N., J. C. Masters, and C. T. Downs. 2010. A population estimate of blue-eyed black lemurs in Ankarafa forest, Sahamalaza-Iles Radama National Park, Madagascar. Folia Primatol. 81: 305-314.
  • Volampeno, M. S. N., J. C. Masters, and C. T. Downs. 2011a. Home range size in the blue-eyed black lemur (Eulemur flavifrons): A comparison between dry and wet seasons. Mamm. Biol. 76(2): 157–164.
  • Blair, M. E., E. J. Sterling, M. Dusch, C. J. Raxworthy, and R. G. Pearson. 2013. Ecological divergence and speciation between lemur (Eulemur) sister species in Madagascar. J. Evolution. Biol. 26(8): 1790–1801.
  • Randriatahina, G. H., and J. J. Roeder. 2013. Group size, composition and stability in a wild population of blue-eyed black lemurs (Eulemur flavifrons) at Ankarafa, Sahamalaza National Park. In J. Masters, M. Gamba, & F. Génin (Eds.), Leaping Ahead (pp. 127–136). Springer New York.
  • Schwitzer, N., D. Clough, H. Zahner, W. Kaumanns, P. Kappeler, and C. Schwitzer. 2010. Parasite prevalence in blue-eyed black lemurs (Eulemur flavifrons) in differently degraded forest fragments. Endang. Species Res. 12: 215-225.
  • Seiler, M., M. Holderied, and C. Schwitzer. 2013. Effects of habitat degradation on sleeping site choice and use in Sahamalaza sportive lemurs. Int. J. Primatol. 34: 260-280.

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Distribution

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The majority of E. flavifrons populations reside on the Sahamalaza Peninsula and adjacent land (Schwitzer et al., 2012 and Volampeno et al., 2013), in the primary and secondary forests (Schwitzer et al., 2012) of northwestern Madagascar (Schwitzer et al. 2007b; Schwitzer et al. 2012; Volampeno et al., 2010). A typical primary forest contains tall, densely packed trees with high species diversity whereas a secondary forest is typically much less densely forested as a result of logging, with less diversity (Schwitzer et al., 2007b). A 2700 Km2 zone comprises most of the habitat of E. flavifrons. This zone includes the Radama National Park, a dry, deciduous forest within the Ankarafa Forest. Most individuals reside in the park (Schwitzer et al. 2012 and Volampeno et al., 2013). The species’ range is almost surrounded by waterways, including the Maevarano, Andranomalaza, and Sandrakota Rivers (Schwitzer et al., 2012). This distribution of E. flavifrons is the smallest among all Eulemur species (Volampeno et al., 2013).

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Brief Summary

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Eulemur macaco has two subspecies: Eulemur macaco macaco (E. macaco) and Eulemur macaco flavifrons (E. flavifrons) (Rabarivola et al., 1991). E. flavifrons is part of the Lemuridae family. The species has two common names: the blue-eyed black lemur and Sclater's lemur; but blue-eyed black lemur is more often used. The medium-sized, 2 Kg species (Schwitzer et al., 2007b) was first described in 1867, by Gray (Mittermieier et al., 2008) and shortly after it was described, it was lost for about 100 years. In 1985, the species was rediscovered (Volampeno et al., 2011a). This species of Eulemur is of interest because it is known to be one of the least-studied species in the genus (Volampeno et al., 2010). Endemic to the north-western part of Madagascar (Schwitzer et al. 2007b; Schwitzer et al. 2012; Volampeno et al., 2010), habitat destruction reduces the species’ success. Logging, slash-and-burn agriculture, forest fragmentation and erosion after deforestation are all forms of habitat destruction that affect the species. Hunting and trapping also reduce the species’ success (Schwitzer et al., 2012; Volampeno et al., 2010; Volampeno et al., 2013). These direct and indirect human effects have decreased the population size of the Eulemur flavifrons species, placing them on the list of critically endangered species (IUCN 2014).

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Behavior

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E. flavifrons is known to have a bi-modal activity pattern, where they are active during both the day and night, with most activity occurring at dusk and dawn (Schwitzer et al., 2012 & Schwitzer et al., 2007a). This activity pattern makes the species cathemeral, which means they distribute their activity over the 24-hour day (Schwitzer et al., 2007a). They are known to forage and feed for about 32% of their day (Goodchild and Schwitzer), which is necessary due to their previously discussed, low-energy diet (Goodchild and Schwitzer, 2008; Volampeno et al., 2011a). A study by Schwitzer et al. (2012) showed that the lemurs’ total daily activity was higher in the secondary forest fragments compared to the primary forest fragments (Schwitzer et al., 2012), which could be due to lower food availability and wider home ranges in the secondary forests. It is also known that during the windy winter season in Madagascar, E. flavifrons has decreased activity on the ground and increased residence in the high trees. In the dry, summer season, the lemurs have an increased activity overall, because they must migrate in search of food (Schwitzer et al., 2007a). Females are known to be dominant in E. flavifrons populations (Digby and Kahlenberg, 2002; Archie and Digby, 1999). When the sexes interact, Digby and Kahlenberg (2002) showed that 99% of the interactions led to the males being subordinate and submissive to the females. The females would use aggression in order to control the males’ actions. All of their seven test groups showed the same results. The females were also aggressive at more times of day than just feeding time, suggesting that they show dominance over all types of interactions (Digby and Kahlenberg, 2002). Juvenile dominance is also present in this species. A study collected data for male offspring that were born to a dominant female and a female offspring that were born to a subordinate female. The study looked at the interactions between the juveniles and other individuals in the population. They found that maternal rank determined whether the juvenile would be dominant in conflicts. Offspring born to the dominant female had a dominant adult to help them in altercations, which made them dominant over others. They reached this conclusion because the juvenile female born to the subordinate mother was more submissive to other individuals in the group compared to the juvenile males with dominant mothers (Archie and Digby, 1999). Behavior between groups is interesting for this species. Each group is composed of about 8 individuals, sometimes a few more or a few less (Randriatahina and Roeder, 2013 & Volampeno et al., 2010). These populations are very stable, with no more than 3 adult females and 2-5 males in one group at a time. Encounters between groups occur at feeding and resting sites, with each group remaining intact, without switching or wandering into another group. Females act antagonistically toward other groups, exerting their dominance (Randriatahina and Roeder, 2013). More research needs to be done in this area to determine if the females limit populations of lemurs from feeding and migrating. If some populations or individuals are not thriving because of females, then management of the populations could include food distribution to see that all individuals are able to eat. E. flavifrons was included in a study of cognitive ability and the similarity between 3 lemur species, old world monkeys, and humans. The study showed that there is conservation among the primate order of the approximate number system, which was used to indicate cognitive ability. The study mentioned that lemurs do not demonstrate deception, or advanced visual communication, or coalition within groups. However, lemurs are capable of transitive learning (understanding that if A>B and B>C, then A<C; Bryson and Leong, 2007), object displacement, and learning contingency (stimulus-based, associative behaviors similar to Pavlovian conditioning; Penn and Povinelli, 2007) when participating in a food-rewarding task (Jones et al., 2012). This study relates to their small group size and feeding habits, indicating that they understand the numerical availability of food (Randriatahina and Roeder, 2013 & Volampeno et al., 2010).

References

  • Archie, E. A. and L. J. Digby. 1999. Juvenile dominance in Eulemur macaco flavifrons: The influence of sex and maternal rank. Folia Primatol. 70(5): 277–81.
  • Bryson, J. J. & J. C. S. Leong. 2007. Primate errors in transitive ‘inference’: a two-tier learning model. Anim Cogn. 10: 1–15
  • Digby, L. J. and S. M. Kahlenberg. 2002. Female dominance in blue-eyed black lemurs (Eulemur macaco flavifrons). Primates 43(3): 191–9.
  • Jones, S. M., J. Pearson, N. K. DeWind, D. Paulsen, A. -M. Tenekedjieva, and E. M. Brannon. 2014. Lemurs and macaques show similar numerical sensitivity. Anim. Cogn. 17(3): 503–515.
  • Penn, D. C. & D. J. Povinelli. 2007. Causal cognition in human and nonhuman animals: a comparative, critical review. Annu. Rev. Psychol. 58: 97–118.
  • Schwitzer, N., Kaumanns, W., P. C. Seitz, and C. Schwitzer. 2007a. Cathemeral activity patterns of the blue-eyed black lemur Eulemur macaco flavifrons in intact and degraded forest fragments. Endang. Species Res. 3: 239-247.

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Life Expectancy

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The life span for E. flavifrons is about 27 years (Volampeno et al., 2011b), and the oldest individual at this time is 21 years old (Volsampeno et al., 2011b).

Reference

Volampeno, M. S. N., J. C. Masters, and C. T. Downs. 2011b. Life history traits, maternal behavior, and infant development of blue-eyed black lemurs (Eulemur flavifrons). Am. J. Primatol. 73: 474-484.

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Management

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As of June 2007, the species habitat was protected in some areas, including the Sahamalaza Peninsula and some surrounding forested areas in the north and the east. Being a UNESCO Biosphere Reserve, the Sahamalaza Peninsula also gives extra protection to the lemurs. There is also a chain of European zoos that are trying to conserve lemur populations by improving and protecting the lemur habitat that is left. A field station in Sahamalaza is also maintained by the group of European zoos, and it is dedicated to current research on lemurs and conservation of the remaining populations (Schwitzer et al., 2012).

Often, parasite levels are used to define stress levels of populations. E. flavifrons was determined to have intestinal parasites in the primary forest fragments and a higher number in the secondary forest fragments. These parasites did not indicate adverse effects on the health of the species, but the higher intestinal parasites in the lemurs that occupied the older, secondary forests could indicate that the environment in the secondary forests is less appropriate for this species (Schwitzer et al., 2012). Forest fragments can be selected for protection based on lemur parasite load. Forest fragments with many parasites can also indicate which populations of the species are especially in need of management (Schwitzer et al., 2012).

As primary forests have been shown to have the highest occupancy of this species, they should be a priority for protection (Schweitzer et al., 2007b). The use of secondary forests as buffer zones and corridors between primary forests would create an ideal dispersal/migration environment for the lemurs. The protection of trees is also a good mechanism to save the lemurs so they will be able to find protection and food (Volampeno et al., 2013). If more research shows that particular primary or secondary fragments of land are more valuable than others, those areas of the habitat should be first priorities for protection and management.

In order to learn more about the species, there needs to be long-term studies in the secondary forests that Eulemur flavifrons occupies (Schwitzer et al., 2007b). If some individuals are kept in captivity, they can be used to appropriately contribute to reintroduction and reinforcement projects (Schwitzer et al., 2013). However, it has been determined that holding lemurs in captivity is not the best option for conservation because they become obese in captive environments, which threatens their health (Goodchild and Schwitzer, 2008). If individuals are to be held captive for research and observation, a proper diet is necessary in order to avoid obesity.

More research is necessary to determine the correct ways in which lemurs need to be managed and conserved. It would be ideal to create a buffer zone around their geographic range to decrease human destruction of their already-fragmented habitat.

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Migration

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E. flavifrons is forced to relocate in order to find food at different times of the year. For example, during the dry season, the food and water shortages causes individuals to travel between primary and secondary forests and between different primary forests in search of food (Volampeno et al., 2011a). During the wet season, however, food is very plentiful. The fruiting trees meet the needs of the lemurs and they will remain in or travel to primary forest fragments to have a more widespread selection of trees to sleep and eat in (Volampeno et al., 2011a).

E. flavifrons could benefit from more research in this area. If the migration of the species for food is causing them to be more vulnerable to threats, then there could be preventative measures taken in order to decrease the need for movement and/or create a way to move between areas safely.

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Molecular Biology

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The molecular difference between E. flavifrons and E. macaco is about 70 base-pairs. According to a genetic analysis, subpopulations of E. flavifrons have been shown to not have a significant difference in their DNA (Fausser et al., 2000). This means that the existing genetic diversity within E. flavifrons is minimal, so it needs to be conserved. This is especially true because E. flavifrons, as of 2000, was not included in many reserves like E. macaco was (Fausser et al., 2000).

Reference

Fausser, J. L., C. Rabarivola, B. Meier, T. Hahn, and Y. Rumpler. 2000. Genetic comparison between different populations of Eulemur macaco flavifrons in northwest Madagascar using RAPD markers. Am. J. Primatol. 51(4): 249–255.

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Morphology

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E. flavifrons is distinguishable from its close genetic relative, E. macaco by its beard, sideburns (Ravarivola et al., 1991), non-tufted ears, and grey-blue eyes (Mittermeier et al., 2008, Rabarivola et al., 1991, Volampeno et al., 2011a). There is also a missing flexion-fold in the palm of E. flavifrons when compared to E. macaco (Rabarivola et al., 1991). E. flavifrons females tend to have a white forehead and lighter fur color than E. macaco (Mittermeier et al., 2008). Females and males experience sexual dichromatism, similar to all but one species of lemur (Mittermeier et al., 2008 and Volampeno et al., 2011b). To have sexual dichromatism, females and males must differ in color. In E. flavifrons, females are reddish-orange whereas males are entirely black (Volampeno et al., 2011b), though they are both medium-sized (2 Kg; Schwitzer et al., 2007b).

E. flavifrons has a flexible visual system that functions in both dim and bright light, confirming that they have both rods and cones within the retina of the eye. More active lemurs have higher visual acuity (Veilleux and Kirk, 2009). E. macaco does not have acute color vision, as it is a prosimian, a primate. This feature is indicated by the way in which the species chooses its foods; bright-colored fruits are not more appealing than dull-colored fruits (Birkinshaw, 2001). These findings are likely applicable to E. flavifrons as it is a subspecies of E. macaco, and also a prosimian.

References

  • Rabarivola, C., D. Meyers, and Y. Rumpler. 1991. Distribution and morphological characters of intermediate forms between the black lemur (Eulemur macaco macaco) and the Sclater’s lemur (E. m. flavifrons). Primates 32(2): 269–273.
  • Birkinshaw, C. 2001. Fruit characteristics of species dispersed by the black lemur (Eulemur macaco) in the Lokobe Forest, Madagascar. Biotropica 33(3): 478-486.
  • Mittermeier, R. A., J. U. Ganzhorn, W. R. Konstant, K. Glander, I. Tattersall, C. P. Groves., A. B. Rylands, A. Hapke, J. Ratsimbazafy, M. I. Mayor, E. E. Louis Jr., Y. Rumpler, C. Schwitzer, & R. M. Rasoloarison. 2008.
  • Lemur diversity in Madagascar. Int. J. Primatol. 29: 1607-1656.
  • Veilleux, C. C., and E. C. Kirk. 2009. Visual acuity in the cathemeral strepsirrhine Eulemur macaco flavifrons. Am. J. Primatol. 71(4): 343–352.
  • Volampeno, M. S. N., J. C. Masters, and C. T. Downs. 2011b. Life history traits, maternal behavior, and infant development of blue-eyed black lemurs (Eulemur flavifrons). Am. J. Primatol. 73: 474-484.

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Population

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It is documented that this species of lemur has the smallest population remaining within the genus. There are about 450-2,300 individuals total, with 24 individuals per km2 as the mean density in 8 different forest fragments. In the Ankarafa forest, there are about 97 individuals per km2 (Schwitzer et al., 2012). The captive population contained 75 individuals as of 2013 (Schwitzer et al., 2013).

References

  • Schwitzer, C., T. King, E. Robsomanitrandrasana, C. Chamberlan, and T. Rasolofoharivelo. 2013. Integrating ex situ and in situ conservation of lemurs. Lemurs of Madagascar: A Strategy for their Conservation 2013–2016. Bristol, UK: IUCN SSC Primate Specialist Group, Bristol Conservation and Science Foundation, and Conservation International. 146-152.
  • Schwitzer, C., G. G. Randriatahina, and S. Volampeno. 2012. Sclater’s black lemur or blue-eyed black lemur, Eulemur flavifrons (Gray 1867). Madagascar.

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Reproduction

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E. flavifrons individuals mate and give birth during the dry season, in which there is decreased food availability. Mating and birth may prevent individuals from foraging and moving in order to find food. This is a problem because less food is available close to home during the dry season, and more traveling is needed to find food (Volampeno et al., 2011a; Volampeno et al., 2011b; Volampeno et al., 2013).

At three years of age, females mate and give birth for the first time (Volampeno et al., 2011b). Before mating season, males migrate toward females. After male migration, there is a male-biased sex ratio which may ensure that the females’ fitness will be higher (Randriatahina and Roeder, 2013). After mating occurs, females disperse throughout the habitat from August to September, which is within the birthing season. Gestation lasts approximately 120 days (Volampeno et al., 2011b), and births occur at the end of August to the end of October, during the dry season (Schwitzer et al., 2012 & Volampeno et al., 2011b). Presumably, in order to avoid predators and excessive attention from group members, females have their babies nocturnally.Many infants are killed by predation, mostly at the time of birth (Volampeno et al., 2011b).

A baby lemur is dependent on its mother until it is 4 weeks old. Afterwards, they can move objects and travel short distances. Mothers need food in order to feed the infants, especially because they are lactating. Birth in the dry season complicates the search for food, as it requires a lot more energy, so instead, the females will rest when they are caring for their baby, instead of traveling far for food, to save energy (Volampeno et al., 2011b). Babies start to explore their environment when they are about 7 weeks old, and are independent at 10 weeks old (Schwitzer et al., 2012; Volampeno et al., 2011b). Females care for their infants by feeding and grooming them until the baby is old enough to do so on its own (Archie and Digby 1999; Volampeno et al., 2011b).

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Threats

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The largest threats to the Eulemur flavifrons population are due to human activities. Humans cause forest destruction and fragmentation, which is usually owed to slash-and-burn agriculture and logging. Trapping and hunting are also a problem (Schwitzer et al., 2012). Currently, 80-90% of the forests that used to exist on Madagascar have been destroyed (Volampeno et al., 2010). A major cause of the destruction of these forests is erosion of the land after logging (Volampeno et al., 2010). There has been a decrease in forest cover from 12.5% to 2.8% in a 40-year time span inside the primary forests, where the lemurs tend to be found more frequently (Volampeno et al., 2013). This reduction in cover means that the lemurs will be more likely to be preyed upon because they are unable to find coverage during sleeping and eating. Since the primary forest fragments are considered the more favorable environment for the species due to the species richness of the fragments, including the presence of fruiting trees and trees for sleeping, human involvement and damage in this ecosystem will reduce the survival of lemur individuals (Schwitzer et al., 2007b).

Reference

Volampeno, M. S. N., G. Randriatahina, and C. T. Downs. 2013. Structure and Composition of Ankarafa Forest, Sahamalaza-Iles Radama National Park, Madagascar: implications for the frugivorous endemic blue-eyed black lemur (Eulemur flavifrons). S. Afr. J. Wildl. Res. 43(2): 91–102.

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Trophic Structure

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E. flavifrons has a diet that mostly consists of fruits, leaves, and flowers (Goodchild and Schwitzer, 2008; Schwitzer et al., 2007b; Volampeno et al., 2011a). In a one-year study, they ate approximately 72 species of plants in the wild. About half of their diet was fruits and the other half was leaves and flowers (Schwitzer, 2012). Volampeno et al. (2012) confirmed that during the wet season, fruit was a large part of the lemurs’ diets, with Magnifer indica as the most-consumed species, aiding in the dispersal of its seeds (Volampeno et al., 2012). Given that E. flavifrons mainly eats fruits and leaves, it seems to be able to survive on a low-energy diet, with a basal metabolic rate that is about 50% lower than most other placental mammals of the same size (Goodchild and Schwitzer, 2008; Volampeno et al., 2011a). E. flavifrons has also been shown to eat small amounts of insects, plant exudates, and fungi (Schwitzer, 2012).

During the day and night, E. flavifrons individuals have to protect themselves from potential predators such as hawks, buzzards, and tree boas. Coverage by trees is essential to protect the lemurs from being preyed upon (Seiler et al., 2013).

References

  • IUCN. 2014. The IUCN Red List of Threatened Species. Version 2014.3. . Downloaded on 17 November 2014.
  • Goodchild, S. and C. Schwitzer. 2008. The problem of obesity in captive lemurs. International Zoo News. 55(6): 353-357.

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Black lemur

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The black lemur (Eulemur macaco) is a species of lemur from the family Lemuridae. Like all lemurs, it is endemic to Madagascar. Originally, the species was thought to have two subspecies,[4] Eulemur macaco macaco and Eulemur macaco flavifrons, both of which were elevated to species status by Mittermeier et al. in 2008 to Eulemur macaco and Eulemur flavifrons respectively.[4] The most startling difference between the two species is the eye colour; Eulemur flavifrons, the blue-eyed black lemur, has blue eyes, while Eulemur macaco, the black lemur, has brown or orange eyes, and also has ear tufts.[5][6]

Both species live in northwest Madagascar. The black lemur occurs in moist forests in the Sambirano region of Madagascar and on nearby islands.[5] The blue-eyed black lemur is restricted to the Sahamalaza Peninsula and adjacent forests.[5] There are reports of the two species hybridizing where their ranges overlap in Manongarivo Special Reserve.[7]

Physical description

Skeleton

The black lemur is between 90 and 110 centimeters in length, of which 51–65 centimeters are tail.[5] Weight typically ranges between 1.8 and 2.0 kilograms.[5] The black lemur displays sexual dimorphism in coloration. Males have black or dark chocolate fur, while females' fur is of a lighter brown color, generally medium brown, chestnut brown or even orange-brown.[5] Males have large black ear tufts, while females have large white ear tufts.

The only other Eulemur species that occurs within the range of the black lemur is the common brown lemur, which overlaps with the black lemur at the extreme southern and eastern edge of its range,[5] and the red-bellied lemur, on the Tsaratanana Massif.[7] The common brown lemur and the red-bellied lemur have different coloration and do not show the extreme sexual dimorphism of the black lemur, making confusion between the black lemur and the other species unlikely.[5][7]

Diet

Male, brown-eyed, Black lemur eating, at Nosy Komba

The black lemur primarily eats fruit,[8] which makes up an estimated 78% of its diet.[9] The ripeness of this fruit is vital to the lemur's diet.[9] Other foods eaten include flowers, leaves, fungi, some invertebrates and, especially during the dry season, nectar.[5]

Behavior

The black lemur lives in both primary and secondary forest.[8] It is active both during the day and at night.[5] It forages in both the upper and middle canopy, especially at night, and during the day it also forages in the understory.[5] In degraded habitats, it also forages on the ground and may even eat soil.[5]

The black lemur lives in groups of 2 to 15 members, including approximately equal numbers of males and females.[5] Average group size is about 10 members,[10] although the average may be smaller for E. m. flavifrons.[5] Females are dominant over males, although intragroup fighting is rare.[8]

Groups have home ranges of about 3.5 to 7 hectares.[10] Ranges overlap considerably, and population density can reach 200 individuals per square kilometer.[8]

Black lemurs also have a habit of picking up and biting at toxic millipedes. The toxins are usually not fatal to the lemurs and they try to stimulate the millipede to release its toxins in self-defence. Once this is achieved the black lemur will rub the millipede around its body to get the toxins on its fur. It is believed that they do this to help repel insects with the millipedes poison, though the toxins also appear to have pleasurable psychoactive effects on the lemurs.[11][12]

Reproduction

Mating usually takes place in April and May.[5] During mating season, antagonism between males increases, and males sometimes roam between groups.[8] After a gestation of about 125 days, a single infant is usually born between late August and early October.[5] Females typically give birth for the first time at two years of age.[8]

Gallery

References

  1. ^ a b Andriantsimanarilafy, R.R.; Borgerson, C.; Clarke, T.; Colquhoun, I.C.; Cotton, A.; Donati, G.; Eppley, T.M.; Heriniaina, R.; Irwin, M.; Johnson, S.; Mittermeier, R.A.; Patel, E.; Ralainasolo, F.B.; Randrianasolo, H.; Randriatahina, G.; Ratsimbazafy, J.; Ravaloharimanitra, M.; Razafindramanana, J.; Schwitzer, C.; Stone, E.; Volampeno, S.; Wright, P. (2020). "Eulemur macaco". IUCN Red List of Threatened Species. 2020: e.T8212A182235113. doi:10.2305/IUCN.UK.2020-3.RLTS.T8212A182235113.en. Retrieved 19 November 2021.
  2. ^ "Checklist of CITES Species". CITES. UNEP-WCMC. Retrieved 18 March 2015.
  3. ^ Groves, C. P. (2005). Wilson, D. E.; Reeder, D. M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Baltimore: Johns Hopkins University Press. p. 115. ISBN 0-801-88221-4. OCLC 62265494.
  4. ^ a b Mittermeier, R. A.; et al. (2008). "Lemur Diversity in Madagascar" (PDF). International Journal of Primatology. 29 (6): 1607–1656. doi:10.1007/s10764-008-9317-y. hdl:10161/6237. S2CID 17614597.
  5. ^ a b c d e f g h i j k l m n o p Nick Garbutt (2007). Mammals of Madagascar. pp. 166–170. ISBN 978-0-300-12550-4.
  6. ^ Noel Rowe (1996). The Pictorial Guide to the Living Primates. pp. 41–42. ISBN 0-9648825-0-7.
  7. ^ a b c Russell Mittermeier; et al. (2006). Lemurs of Madagascar (Second ed.). pp. 288–293. ISBN 1-881173-88-7.
  8. ^ a b c d e f Robert W. Sussman (1999). Primate Ecology and Social Structure Volume 1: Lorises, Lemurs and Tarsiers. pp. 186–187. ISBN 0-536-02256-9.
  9. ^ a b Rakotosamimanana, Berthe; Hanta Rasamimanana (1999). New Directions in Lemur Studies. Springer. ISBN 0-306-46187-0.
  10. ^ a b Lisa Gould & Michelle Sauther (2007). "Lemuriformes". In Christina J. Campbell; Agustin Fuentes; Katherine C. MacKinnon; Melissa Panger & Simon K. Bearder (eds.). Primates in Perspective. p. 53. ISBN 978-0-19-517133-4.
  11. ^ Downer, John (25 April 2002). "Peculiar Potions: Narcotic insecticide". Weird Nature. Series 1. Episode 4. John Downer Productions. BBC Four. Retrieved 22 March 2023.
  12. ^ Banerji, Urvija (29 January 2016). "Lemurs Get High on Their Millipede Supply". Atlas Obscura. Retrieved 22 March 2023.

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Black lemur: Brief Summary

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The black lemur (Eulemur macaco) is a species of lemur from the family Lemuridae. Like all lemurs, it is endemic to Madagascar. Originally, the species was thought to have two subspecies, Eulemur macaco macaco and Eulemur macaco flavifrons, both of which were elevated to species status by Mittermeier et al. in 2008 to Eulemur macaco and Eulemur flavifrons respectively. The most startling difference between the two species is the eye colour; Eulemur flavifrons, the blue-eyed black lemur, has blue eyes, while Eulemur macaco, the black lemur, has brown or orange eyes, and also has ear tufts.

Both species live in northwest Madagascar. The black lemur occurs in moist forests in the Sambirano region of Madagascar and on nearby islands. The blue-eyed black lemur is restricted to the Sahamalaza Peninsula and adjacent forests. There are reports of the two species hybridizing where their ranges overlap in Manongarivo Special Reserve.

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