Animal Diversity Web

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)

• Biogeographic Regions
• ethiopian
  • native

• Other Geographic Terms
• island endemic

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)

• Habitat Regions
• tropical
• terrestrial

• Terrestrial Biomes
• rainforest

• Other Habitat Features
• agricultural
• riparian

• 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)

• Other Physical Features
• endothermic
• bilateral symmetry

• 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)

• Mating System
• polygynandrous (promiscuous)

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)

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

• 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)

• Key Behaviors
• arboreal
• diurnal
• nocturnal
• crepuscular
• motile
• social
• dominance hierarchies

• 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)

• Communication Channels
• visual
• tactile
• acoustic
• chemical

• Other Communication Modes
• scent marks

• Perception Channels
• visual
• tactile
• acoustic
• chemical

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)

• Primary Diet
• herbivore
  • frugivore

• 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)

• Known Predators

  • harrier hawks (Polyboroides typus)
  • humans (Homo sapiens)
  • fossas (Cryptoprocta ferox)

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

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)

• Positive Impacts
• pet trade
• food
• ecotourism
• pollinates crops

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)

• IUCN Red List

  Vulnerable
  More information

• IUCN Red List

  Vulnerable
  More information

• US Federal List

  Endangered

• CITES

  Appendix I

Con­trib­u­tors

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

Ref­er­ences

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