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Eulemur macacoblack lemur

By Lauren Gravatt

Geographic Range

Black lemurs are restricted to the areas of northwestern Madagascar surrounding the Ampasindava Peninsula and Narinda Bay. Black lemurs also inhabit the close neighboring islands of Nosy Be and Nosy Komba. Their range covers a total fragmented area of approximately 11,740 km2. (Andriaholinirina, et al., 2014; Nowak and Walker, 1999)

Habitat

The habitat of Eulemur macaco is made up of Madagascar’s wet evergreen, dry deciduous and riverine forests. These canopy dwellers occupy primary and secondary rainforests as well as timber and food crop plantations. Black lemurs live at elevations ranging from sea level up to 1600 m. (Andriaholinirina, et al., 2014; Fleagle, et al., 1999)

  • Range elevation
    0 to 1600 m
    0.00 to 5249.34 ft

Physical Description

Eulemur macaco can range from 1.5 to 3 kg with an average body mass of 1.8 kg in the wild and 2.5 kg in captivity. From head to rump, they are approximately 30 to 50 cm and their tail extends another 40 to 60 cm.

Black lemurs display sexual dichromatism in their coat color. The males are solid with a dark chocolate brown to black pelage while females have chestnut to golden brown bodies with lighter ventral surfaces and black muzzles and feet. Black lemurs have forward facing eyes and an elongated rostrum with a rhinarium. Eulemur macaco has brown or orange eyes that help to distinguish it from its relative the blue-eyed black lemur. Eulemur macaco has a soft, relatively lengthy coat and a long bushy tail. Both sexes have tufts of hair that frame their face from the throat to just above the ears. This ruffed hair is black in males and white to off-white in females.

Eulemur macaco has four toes with an opposable pollex on each hand and each hind foot has four toes with an opposable hallux. All of the digits have nails except the second digit on the hind foot which instead has a grooming claw. The hind limbs of E. macaco are longer than the forelimbs.

According to Nutrient Requirements of Nonhuman Primates: Second Revised Edition, in general, the basal metabolic rate of a 2.5 kg lemur is 110 kcal/kg. Thus, for a 1.8 kg black lemur it would be approximately 198 kcal. During gestation, energetic needs increase by about 2.5 to 3%, and basal metabolic rate increases to 256 to 301 kcal*BWkg.75, so approximately 397.8 to 467.75 kcal for the average 1.8 kg black lemur.

The general dental formula for members of the family Lemuridae is 2/2, 1/1, 3/3, 3/3 = 36. The lower incisors and canines of lemurs project forward to form a tooth comb used for grooming. ("Nutrient Requirements of Nonhuman Primates", 2003; AZA Prosimian Taxon Advisory Group, 2013; Fleagle, et al., 1999; Nowak and Walker, 1999; Patton, 2008)

  • Sexual Dimorphism
  • sexes colored or patterned differently
  • Range mass
    1.5 to 3 kg
    3.30 to 6.61 lb
  • Average mass
    1.8 kg
    3.96 lb
  • Range length
    70 to 110 cm
    27.56 to 43.31 in
  • Range basal metabolic rate
    165 kcal to 330 kcal cm3.O2/g/hr
  • Average basal metabolic rate
    198 kcal cm3.O2/g/hr

Reproduction

Eulemur macaco has a complex promiscuous mating system. Male mating behavior involves scent marking females, clasping them, and ano-genital grooming. Male black lemurs display roaming behavior during the breeding season by leaving their social group to follow females from other groups. Roaming males have been seen engaging in mating behavior such as scent marking and ano-genital grooming with the females from these neighboring groups.

Dominance doesn’t seem to influence the roaming behavior, as both subordinate and dominant males have been witnessed roaming. Dominant males have been seen guarding females in the group from subordinate and foreign males. When the dominant male left to roam, subordinate males were able to copulate with the females he had been guarding.

Female choice seems to be a major theme in the mating system of E. macaco. Females have the choice to mate with dominant, subordinate or roaming males, and also the authority to reject males. Females can deter unwanted males with agonistic techniques and by guarding their genitals with their tail.

Although intergroup copulation has not been witnessed in black lemurs, the roaming behavior observed by males along with the observations of ring-tailed lemurs, the ring-tailed lemur, breeding with members of different social groups, suggests that intergroup reproduction is a possibility in E. macaco. (Colquhoun, 1997)

Eulemur macaco is iteroparous with a breeding interval of one year. Black lemurs breed between April and May and give birth to altricial young from late August to November. The gestation period is 125 to 127 days. In general, females have one offspring per year, though in captivity, twinning rates are reported at 21%

Female black lemurs give birth to altritial young that weigh approximately 66g, which is about 5.4% of the mother’s body mass. Infant weaning occurs between 5 and 7 months of age in the wild, and 3 to 4 months in captivity. Juvenile males and females could disperse to new groups between one and two years of age. Male and female black lemurs reach sexual maturity at approximately 548 days of age. (AZA Prosimian Taxon Advisory Group, 2013; Andriaholinirina, et al., 2014; Bayart and Simmen, 2005; Colquhoun, 1997; Nowak and Walker, 1999)

  • Breeding interval
    Black lemurs breed once yearly
  • Breeding season
    April to May
  • Range number of offspring
    1 to 2
  • Average number of offspring
    1.04
  • Average number of offspring
    1
    AnAge
  • Range gestation period
    120 to 129 days
  • Average gestation period
    125 days
  • Range weaning age
    3 to 7 months
  • Range time to independence
    1 to 2 years
  • Average age at sexual or reproductive maturity (female)
    548 days
  • Average age at sexual or reproductive maturity (female)
    Sex: female
    548 days
    AnAge
  • Average age at sexual or reproductive maturity (male)
    548 days
  • Average age at sexual or reproductive maturity (male)
    Sex: male
    548 days
    AnAge

Black lemurs have constant contact with their mother for the first month and suckle the majority of the time. For the first three weeks of life, the infant clings to its mother’s belly while she locomotes and when not moving the mother leans over the infant covering it completely. After three weeks, the infant begins to ride on its mother’s back. The first complete separation of mother and offspring occurs around 40-45 days of age. As the infant grows older, it begins to spend more time away from its mother, though it may still be nursing at 5-6 months of age.

Females keep all social group members away from the newborns after giving birth. Male parental investment has not been well documented, though it is noted that the males do not go anywhere near a mother and newborn until the newborn is at least several days old. (Colquhoun, 1997; Jolly, 2005; Tilden and Oftedal, 1997)

  • Parental Investment
  • altricial
  • female parental care
  • pre-weaning/fledging
    • provisioning
      • female
    • protecting
      • female
  • extended period of juvenile learning

Lifespan/Longevity

The lifespan of black lemurs in the wild has not been well documented, though it is expected to be around 15 to 20 years. In captivity, on average, black lemurs live approximately 20 to 25 years. The longest known lifespan of Eulemur macaco was 36.2 years, though a hybrid of blue-eyed black lemur Eulemur flavifrons and Eulemur macaco lived to 39 years in captivity. ("AnAge Entry for Eulemur macaco", 2014; "Black Lemur (Eulemur macaco)", 2014; Simmen, et al., 2007)

  • Range lifespan
    Status: captivity
    36.2 (high) years
  • Typical lifespan
    Status: wild
    15 to 20 years
  • Typical lifespan
    Status: captivity
    20 to 25 years

Behavior

Black lemurs live in social groups of 4 to 15 individuals consisting of multiple males and females. The average group size is 7, and in general there is approximately a 1:1 distribution of males and females. Females are the dominant sex within social groups of black lemurs and get priority for resources such as food and water. The core of groups is made of mature females. While there is some evidence suggesting a strong male-female bond within groups, frequent emigration of males and roaming mating behavior may suggest otherwise.

Eulemur macaco uses quadrupedal arboreal locomotion and arboreal leaping locomotion from branch to branch or tree to tree. On the ground they are quadrupedal with bouts of bipedal locomotion. The amount of time black lemurs allocate to traveling and resting is approximately equal among groups and seasons. However, the range and speed of travel is greatly reduced during the dry seasons when resources may be scarcer.

Black lemurs have an average territory size of 3.5 to 7 ha, with a mean home range from 5 to 5.5 ha. Group home ranges tend to overlap considerably and group encounters are not uncommon. During these encounters, physical harm has not been witnessed but territorial displays such as tail swishing and leaping back and forth are common. During mating season, there is an increased amount of intergroup interaction by the roaming males, and during birthing season there is greater intergroup avoidance.

During the mating season black lemurs also spend less time eating and more time finding mates. Black lemur males have increased agonistic interactions toward other males during the breeding season and females have increased agonistic interactions over fruit during lactation. Black lemurs do not tend to show reconciliatory post-conflict behavior, they instead disperse after a conflict.

Eulemur macaco is a cathemeral primate, meaning that it has a 24 hour activity cycle that is neither strictly diurnal nor solely nocturnal. Black lemurs display significant bouts of activity during both diurnal and nocturnal periods. They seem to be the most active from 06.00 h to 08.00 h and from 16.00 h to 18.00 h. E. macaco is relatively inactive during the middle hours of the day. With the exception of the hours between 04.00 h and 06.00 h, nocturnal activity is greater during light moon nights than dark moon nights. It is hypothesized that this could be due to lemur eyes lacking a tapetum lucidum, the structure in the eye that enhances night vision by increasing the amount of light available for photoreceptors. Without this structure, lemurs may have a difficult time seeing on nights without bright moonlight. Nocturnal activity is not linked to the lunar cycle between the hours of 04.00 h and 06.00 h when the sky begins to lighten as the sun nears the horizon, presumably because the lemurs have better vision during this time.

Black lemurs have been documented multiple times antagonizing toxic millipedes in order to rub the toxic secretions on their fur. They gently bite the millipede several times until it secretes the toxins, and then cover their bodies with the toxins. It is presumed that the millipede toxin has insecticidal properties that are beneficial to the lemurs. A side effect of the toxins leaves the lemurs with a “high” sensation. In most cases, the millipede is released relatively unscathed. (Bayart and Simmen, 2005; Colquhoun, 1997; Colquhoun, 1998; Downer and McMenamin, 2014; Lee, 1999; Nowak and Walker, 1999; Olliver, et al., 2004)

  • Range territory size
    35,000 to 70,000 m^2
  • Average territory size
    50,000 to 55,000 m^2

Home Range

Black lemurs have an average territory size of 3.5 to 7 ha, with a mean home range from 5 to 5.5 ha. Group home ranges tend to overlap considerably. (Bayart and Simmen, 2005; Colquhoun, 1997)

Communication and Perception

Black lemurs can communicate by scent marking, vocalizing, and body language. They scent mark with excretions from unique perianal glands. Distinct vocalizations such as grunting, cackling, and screeching can be used to communicate. Body language, touch and facial expression are also important aspects of communication by Eulemur macaco. Females may display their dominance with antagonistic actions such as chasing, cackling or cuffing more subordinate members of the group. ("Black Lemur (Eulemur macaco macaco)", 2014; Bayart and Simmen, 2005; Colquhoun, 1997)

Food Habits

Black lemurs are mostly frugivorous with fruit making up an average amount of 73% of their diet. The other components of their diet include about 13% flowers and nectar, 12% foliage, 2% bark and small amounts of earth and ants. During the dry season, the majority of the diet is made up of Coffea (coffee), Corica papaya (papaya), Dypsis (palm), Anacardium occidentale (cashew), and Cordia myxa (plum). Corica papaya (papaya), Mangifera indica (mango), palm fruit and flowers from Dypsis and flowers of Grewia make up most of the diet during the wet season. During the day, between 05.00 h and 19.00 h Eulemur macaco ingests 420 to 530 g of food, and another 120 g of food during the night.

Eulemur macaco has displayed strong dietary diversity in the wild and in captivity. While there is no evidence of carnivorous behavior in the wild, black lemurs have accepted meat as food in captivity. (Nowak and Walker, 1999; Simmen, et al., 2007)

  • Animal Foods
  • insects
  • Plant Foods
  • leaves
  • wood, bark, or stems
  • seeds, grains, and nuts
  • fruit
  • nectar
  • flowers

Predation

Predators of Eulemur macaco include raptors, humans, and viverrids. Anti-predator adaptations include specific vocalizations, mob behavior, alarm calls, staring, shrieking and tail wagging. In response to terrestrial predators, black lemurs often huff, grunt, tail wag, and can display full mobbing behavior. In response to fossa, E. macaco will stare, grunt, shriek and flee upwards. They will alarm call with screams and whistles and take cover in the lower canopy in response to aerial predators. Mob behavior has been documented against boa constrictors and harrier hawks. If the mob behavior is not successful, they retreat. Black lemurs rely on safety in numbers and alarm call warnings. ("Black Lemur (Eulemur macaco macaco)", 2014; Colquhoun, 1993; Gursky and Nekaris, 2007)

Ecosystem Roles

Black lemurs are thought to play a very important role in seed dispersal of fruit seeds through their feces. It has also been suggested that they may act as pollinators for the flowers on which they are nectivorous.

While there’s no information on the parasites of Eulemur macaco, its close relative, the blue-eyed black lemur Eulmer flavifrons does have parasitic information. In Eulemur flavifrons, about 13% of tested fecal samples contained intestinal parasite eggs, while 55% contained adult intestinal parasites. The parasites include Callistoura and Lemuricola. There were also ear mites present in 33% of tested lemurs. Parasite abundance of Eulemur flavifrons was greater in secondary forests than primary, suggesting that there were greater stresses and hardships in the secondary forests.

Black lemurs have a potentially commensal relationship with the toxic giant African millipede Spirostreptus gigas. The lemurs irritate the millipede in order to use their secretions as insecticide. The millipedes are relatively unharmed and usually survive the encounter. ("Black Lemur (Eulemur macaco macaco)", 2014; Downer and McMenamin, 2014; Schwitzer, et al., 2010)

  • Ecosystem Impact
  • disperses seeds
  • pollinates
Commensal/Parasitic Species
  • Callistoura
  • Lemuricola

Economic Importance for Humans: Positive

Black lemurs are often hunted as food and sold as pets for natives in Madagascar. In general,members of the family Lemuridae tend attract tourists to the island of Madagascar which may help support the local economy. Black lemurs also provide seed dispersal through feces and may help pollinate certain flowers. (Andriaholinirina, et al., 2014)

Economic Importance for Humans: Negative

Eulemur macaco may be considered a crop pest in some areas where fruit or cashews are farmed by locals. (Nowak and Walker, 1999)

  • Negative Impacts
  • crop pest

Conservation Status

The U.S. Federal List considers all lemurs to be endangered, however, the ICUN Red Book lists Eulemur macaco as vulnerable and declining. Black lemurs are listed as an Appendix I species by CITES. Black lemurs are experiencing severe habitat loss and fragmentation. They may also be over exploited by the Malagasy people for food, pets and zoos. Black lemurs are also killed as crop pests. (Andriaholinirina, et al., 2014)

Contributors

Lauren Gravatt (author), University of Alaska Fairbanks, Laura Prugh (editor), University of Washington, Tanya Dewey (editor), University of Michigan-Ann Arbor.

Glossary

Ethiopian

living in sub-Saharan Africa (south of 30 degrees north) and Madagascar.

World Map

acoustic

uses sound to communicate

agricultural

living in landscapes dominated by human agriculture.

altricial

young are born in a relatively underdeveloped state; they are unable to feed or care for themselves or locomote independently for a period of time after birth/hatching. In birds, naked and helpless after hatching.

arboreal

Referring to an animal that lives in trees; tree-climbing.

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

chemical

uses smells or other chemicals to communicate

crepuscular

active at dawn and dusk

diurnal
  1. active during the day, 2. lasting for one day.
dominance hierarchies

ranking system or pecking order among members of a long-term social group, where dominance status affects access to resources or mates

ecotourism

humans benefit economically by promoting tourism that focuses on the appreciation of natural areas or animals. Ecotourism implies that there are existing programs that profit from the appreciation of natural areas or animals.

endothermic

animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.

female parental care

parental care is carried out by females

food

A substance that provides both nutrients and energy to a living thing.

frugivore

an animal that mainly eats fruit

herbivore

An animal that eats mainly plants or parts of plants.

island endemic

animals that live only on an island or set of islands.

iteroparous

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

motile

having the capacity to move from one place to another.

native range

the area in which the animal is naturally found, the region in which it is endemic.

nocturnal

active during the night

pet trade

the business of buying and selling animals for people to keep in their homes as pets.

polygynandrous

the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.

rainforest

rainforests, both temperate and tropical, are dominated by trees often forming a closed canopy with little light reaching the ground. Epiphytes and climbing plants are also abundant. Precipitation is typically not limiting, but may be somewhat seasonal.

riparian

Referring to something living or located adjacent to a waterbody (usually, but not always, a river or stream).

scent marks

communicates by producing scents from special gland(s) and placing them on a surface whether others can smell or taste them

seasonal breeding

breeding is confined to a particular season

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

social

associates with others of its species; forms social groups.

tactile

uses touch to communicate

terrestrial

Living on the ground.

tropical

the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.

visual

uses sight to communicate

viviparous

reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.

References

2014. "AnAge Entry for Eulemur macaco" (On-line). The Animal Aging and Longevity Database. Accessed November 15, 2014 at http://genomics.senescence.info/species/entry.php?species=Eulemur_macaco.

2014. "Black Lemur (Eulemur macaco macaco)" (On-line). Point Defiance Zoo and Aquarium. Accessed November 15, 2014 at http://www.pdza.org/black-lemur/.

2014. "Black Lemur (Eulemur macaco)" (On-line). San Francisco Zoo and Gardens. Accessed November 15, 2014 at http://www.sfzoo.org/explore/animals/mammals/blacklemur.htm.

1976. "Lemurs (Lemuridae entire)" (On-line). U.S. Fish and Wildlife Service: Environmental Conservation Online System. Accessed December 10, 2014 at http://ecos.fws.gov/speciesProfile/profile/speciesProfile.action?spcode=A05P.

National Academy of Sciences. Nutrient Requirements of Nonhuman Primates. SF407.P7 N88. Washington, DC: National Academies Press. 2003.

AZA Prosimian Taxon Advisory Group, 2013. Eulemer Care Manual. Silver Spring, MD: Association of Zoos and Aquariums.

Andriaholinirina, N., A. Baden, M. Blanco, L. Chikhi, A. Cooke, N. Davies, R. Dolch, G. Donati, J. Ganzhorn, C. Golden, L. Groeneveld, A. Hapke, M. Irwin, S. Johnson, P. Kappeler, T. King, R. Lewis, E. Louis, M. Markolf, V. Mass, R. Mittermeier, B. Rakotomanga, J. Rakotonanahary, H. Rakotondrainibe, G. Rakotondratsimba, M. Rakotondratsimba, L. Rakotonirina, F. Ralainasolo, J. Ralison, T. Ramahaleo, H. Randrianasolo, G. Randriatahina, H. Rasamimananana, T. Rasolofoharivelo, S. Rasoloharijaona, F. Ratelolahy, J. Ratsimbazafy, N. Ratsimbazafy, H. Razafindraibe, J. Razafindramanana, N. Rowe, J. Salmona, M. Seiler, S. Volampeno, P. Wright, J. Youssouf, J. Zaonarivelo, A. Zaramody. 2014. "Eulemur macaco" (On-line). IUCN Red List of Threatened Species. Accessed November 16, 2014 at http://www.iucnredlist.org/details/full/8212/0.

Bayart, F., B. Simmen. 2005. Demography, Range Use, and Behavior in Black Lemurs (Eulemur macaco macaco) at Ampasikely, Northwest Madagascar. American Journal of Primatology, 67.3: 299-312.

Colquhoun, I. 1997. A Predictive Socioecological Study of the Black Lemur (Eulemur macaco macaco) in Northwestern Madagascar. Saint Louis, Missouri: Washington University.

Colquhoun, I. 1998. Cathemeral Behavior of Eulemur macaco macaco at Ambato Massif, Madagascar. Folia Primatologica, 69.1: 22-34.

Colquhoun, I. 1993. The Socioecology of Eulemur macaco: A Preliminary Report. Lemur Social Systems and Their Ecological Basis, 31.9: 11-23.

Downer, J., C. McMenamin. 2014. "Peculiar Potions: Narcotic Insecticide" (On-line video). Nature Wildlife. Accessed December 08, 2014 at http://www.bbc.co.uk/nature/life/Lemuridae#p00zd3w2.

Fleagle, J., C. Helmar, J. Reed, K. Reed. 1999. Primate Communities. Cambridge: Cambridge UP.

Gursky, S., K. Nekaris. 2007. Primate Anti-predator Strategies. New York, NY: Springer.

Jolly, A. 2005. "Black Lemur (Eulemur macaco)" (On-line). Wildscreen Arkive. Accessed December 08, 2014 at http://www.arkive.org/black-lemur/eulemur-macaco/.

Lee, P. 1999. Comparative Primate Socioecology. Cambridge, UK: Cambridge UP.

Nowak, R., E. Walker. 1999. Walker's Primates of the World. Baltimore: Johns Hopkins UP.

Olliver, F., D. Samuelson, D. Brooks, P. Lewis, M. Kallberg, A. Komaromy. 2004. Comparative Morphology of the Tapetum Lucidum (among Selected Species). Veterinary Ophthalmolgy, 7.1: 11-22.

Patton, J. 2008. Orders and Families of Living Mammals: A Lab Manual for Mammalogy, BIOL 425. Berkeley: Museum of Vertebrate Zoology, University of California.

Schwitzer, N., D. Clough, H. Zahner, W. Kaumanns, P. Kappeler, C. Schwitzer. 2010. Parasite Prevalence in Blue-eyed Black Lemurs Eulemur flavifrons in Differently Degraded Forest Fragments. Endangered Species Research, 12.3: 215-225.

Simmen, B., F. Bayart, A. Marez, A. Hladik. 2007. Diet, Nutritional Ecology, and Birth Season of Eulemur macaco in an Anthropogenic Forest in Madagascar. International Journal of Primatology, 28.1: 253-266.

Tilden, C., O. Oftedal. 1997. Milk Composition Reflects Pattern of Maternal Care in Prosimian Primates. American Journal of Primatology, 41.3: 195-211.

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