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Varecia variegatavariegated lemur

By Meg Hale

Geographic Range

The black-and-white ruffed lemur, Varecia variegata, is endemic to the eastern rainforest of Madagascar, located in the Indian Ocean. The geographical range of this species includes areas south of the Mananara River in southeastern Madagascar and extends north to the Antainambalana River, northwest of the Bay of Antongil.

Black-and-white ruffed lemurs have been introduced to the Betampona Reserve, located northwest of the major port of Tamatave. They have also been introduced to Nosy Mangabe, which is an island located off the eastern coast of Madagascar. (Baden, et al., 2008; Britt, 1998; Hekkala, et al., 2007)

Habitat

The habitat of the black-and-white ruffed lemur is primary rain forests in eastern Madagascar. These lemurs are arboreal and spend most of their time in the canopy of trees. The elevation of the rain forests in eastern Madagascar can reach 800 meters, they receive 200 centimeters of rain each year. (Louis, Jr, et al., 2005; Wright, 1999)

  • Range elevation
    0 to 800 m
    0.00 to 2624.67 ft

Physical Description

The largest species in their family, the body mass of adult black-and-white ruffed lemurs range from 2.5 kg to 4.8 kg. On average, females weigh more than males, especially during breeding and gestational periods. The female lemurs have notably longer tails than the males. The length of the males is between 436mm and 565mm (average = 493mm). The length of the females is between 469mm and 570mm (average = 516mm).

Male and female black-and-white ruffed lemurs have long hair with black and white markings, each individual with varied amounts of black and white. Their hands, feet, face, and tails are black. They have a white headcrown that is on average 12.2 cm.

At birth, lemurs weigh an average of 87.2 grams and are furred with black and white coloration. (Baden, et al., 2008; Kohler, et al., 2006; Nelson, et al., 2009)

  • Sexual Dimorphism
  • female larger
  • Range mass
    2.5 to 4.8 kg
    5.51 to 10.57 lb
  • Range length
    436 to 570 mm
    17.17 to 22.44 in

Reproduction

Roughly 6 weeks prior to sexual intercourse, the males' testicle size grows and they become more aggressive towards one another. The females' vaginas open to begin estrus so intercourse can occur. Estrus last for about one week out of the year.

Male lemurs approach females by making a whining noise. He will then sniff and lick around the females’ anogenital region. Although the males are submissive towards the females during their estrus phase (due to heightened female aggression), they will still try to mount the female. Females have also been observed to approach a male and slap at him, leading to mounting and intercourse. Female lemurs are polyandrous, meaning they mate with multiple males. (Pereira, et al., 1988; Rasmussen, 1985)

Ruffed lemurs reproduce once a year in the fall or winter. Female lemurs undergo estrus for about a week. Only during this time, sexual intercourse occurs. Lemurs do not engage in intercourse any other time of the year. The gestation period for this species is 117 days. The litter size is 1 to 3 offspring. The average birth mass of a new born lemur is 87.2 grams. Babies are altrical and stay in their original natal nest for 2 to 3 weeks before nest transfer begins. Nest transfers consists of females moving their litter among multiple nests throughout the day to allow for foraging in different locations. These nests were built prior to the birth of the offspring. The babies are weaned from their mothers around 7 to 8 months and become more independent. Female lemurs reach sexual maturity about 604 days after birth. Males reach sexual maturity 608 days after birth. (Baden, et al., 2013; Rasmussen, 1985)

  • Breeding interval
    Once yearly for about a week
  • Breeding season
    Fall or Winter
  • Range number of offspring
    1 to 3
  • Average number of offspring
    2.2
    AnAge
  • Range gestation period
    100 to 135 days
  • Average gestation period
    117 days
  • Range weaning age
    7 to 8 months
  • Average age at sexual or reproductive maturity (female)
    604 days
  • Average age at sexual or reproductive maturity (female)
    Sex: female
    604 days
    AnAge
  • Average age at sexual or reproductive maturity (male)
    608 days
  • Average age at sexual or reproductive maturity (male)
    Sex: male
    608 days
    AnAge

Roughly 3 weeks prior to giving birth, black-and-white ruffed lemurs build up to 15 natal nests in different trees, to care for their offspring. The nests are built 10-20 meters high. Each mother constructs her many nests individually.

Nest transfers begin once the infants reach 2 to 3 weeks of age. The mother carries her infants by mouth to transport them. Nest location changes 1-5 times per day in the first month and increase to 3-7 times per day in the second month. The lemurs move nest to nest for foraging purposes. The mother leaves her infants in the nests for a couple hours at a time while she forages for food. Lemurs with different mothers are not placed in the same nests together.

For the first 6 weeks, the mother is the sole provider for her offspring and most of her time is spent foraging for food and resting with and nursing her young. After 6 weeks, some lemurs begin a method of communal nesting, where most lemurs in the same community help each other to provide for their offspring. The Father lemurs then resume contact with their offspring by grooming and playing with them. Some lemurs only crèche (caring for anothers infant) with relative males and females, while others crèche with unrelated and/or nonbreeding males and females. Alloparental lemurs are the individuals who help crèche specific offspring. These alloparental lemurs groom, play, and travel closely with the infants. They also look after the young while the mother forages for food.

Not all of the lemurs participate in the communal crèche method. The females that decide to raise their young exclusively are called single nesters. Communal nesting contributes to the reproductive success of this species and ensures infant survival rate. Current research on this species is working to explain why certain ruffed lemurs do not engage in communal nesting.

Throughout their childhood, young lemurs begin to display and model the adult behaviors in their community. At age 2 to 3 months, lemurs learn to travel closely behind the adults because they are too large to be carried in the mouth of their mother; at this point they are able to travel up to 50 to 100 meters. When the young reach 7-8 months, they are weaned from their mother and no longer need to continue nursing. Once the adolescents become more independent, the mothers begin to spend less time foraging and caring for their offspring and more time interacting and traveling. The adolescents grow up to become a member of their mothers' social group. (Baden, et al., 2013; Morland, 1990)

  • Parental Investment
  • altricial
  • female parental care
  • pre-hatching/birth
    • provisioning
      • female
  • pre-weaning/fledging
    • provisioning
      • female
    • protecting
      • male
      • female
  • pre-independence
    • provisioning
      • male
      • female
    • protecting
      • male
      • female
  • post-independence association with parents
  • extended period of juvenile learning

Lifespan/Longevity

Black-and-white ruffed lemurs live in the wild and in captivity for an average of 19 years. In the wild, the oldest known lemur lived to be 37 years old. In captivity, the oldest known lemur lived to be 33.4 years old. Female lemurs tend to live a couple years longer than males. (Hakeem, et al., 1996; Kohler, et al., 2006)

Behavior

Black-and-white ruffed lemurs stand quadrupedally with an arch in their back. They hold their tails upward in a question mark shape or in an anterior facing C shape. The lemurs sit hunched over, semi-hunched, and upright. When in the hunched position, the lemurs lean forward, wrap their tails around their body, and hold their head at or below shoulder level, sometimes resting it in their hands. This is their common sleeping position. While sitting in the Semi-hunched position, the lemurs arch their back forward, put their hands flat, lock their elbows, and lean their weight towards their arms. Their head is held vertically above their shoulders. During sunning and relaxation, the lemurs sit in an upright position with most of their weight on their ischium, no arch in their spine, and their legs are either out straight and spread apart or pulled in positioned close to their torso.

Lemurs lay in a supine position flexing their hind legs and extending their arms forward or laterally. They also lay in a prone position on a wide tree branch with their legs and arms relaxed, hanging down. Regularly, these lemurs will lie with their torso twisted causing part of their body to be on its side and the torso in a flat position.

Ruffed lemurs hang bipedally, holding themselves upside down, body extended, with their feet latched onto a tree branch. They also hang with their feet and one hand or with both hands and both feet, but never hang by their hands alone. In the hanging position they groom, eat, play, and relax. They do not hang upside down for longer than 45 seconds.

While walking, ruffed lemurs use all four limbs, point their head downward, and hold their tails upright. Lemurs run by pushing with their hind legs and reaching their arms forward with one arm ahead of the other, called a leading limb. They jump using all four limbs and land on all four limbs. When jumping vertically into a tree, the lemurs push off with all fours and extend their hands, reaching for a branch. Their hands make the first contact with the branch. They travel tree to tree by leaping, moving under branch supports (suspension), and moving across terminal branches (bridging).

Ruffed lemurs forage for food on the forest floor and in trees. They grab their food with their hands or by mouth. While eating, they hold their head vertically to keep the food in between their cheeks and teeth.

Scent-marking is a common way of communicating among ruffed lemurs throughout the year, though more frequent in breeding seasons, and begins around 6 months of age. Females scent-mark by squatting and rubbing their anogenital region back and forth on a substrate. Males sent mark by lowering their body and rubbing their chin, chest, and neck on a surface. Males and females only scent-mark on horizontal surfaces.

Pereira, Seeligson, and Macedonia (1988) report that adolescent lemurs spend 8% of their time feeding, 15% traveling, 9% interacting with their social group, and 55% resting. The adults spend 9% of their time feeding, 18% traveling, 8% interacting with their social group and 45% resting. These animals travel together as a group. Inconsistently, subgroups split away from the main group to forage and rest for extended periods of time. Any member of the social group can initiate and lead traveling to a particular location. There are no social hierarchy behaviors.

The group members sniff each other as a greeting, which often leads to social grooming. Grooming one other begins at the age of 5 months. Self-grooming is more common in lemurs than social grooming throughout the day. (Baden, et al., 2013; Morland, 1990; Pereira, et al., 1988)

Home Range

The lemurs normally occupy 3.5 ha of land with their social groups in their geographical range. The lemurs protect their territory by making threatening calls to other lemur communities trying to encroach in their current territory. (Pereira, et al., 1988)

Communication and Perception

Black-and-white ruffed lemurs have two different types of communicating calls. One of their calls is an inconsistent series of roaring with irregular high pitched squeals throughout their call. This type of communication signals the surrounding species with a warning that there is a predator nearby. This sound is also a call out from the male lemurs when a female lemur has just rejected his effort to mate with her. The other type of communicating call is a series of short chucking sounds, which is a territorial spacing signal to other groups of lemurs. Evidence has shown that this same sound is their way of communicating about a potential predator. Gender does not play a role in the pitch, pulse rate, and frequency ranges during communication.

Lemurs perceive their environment through visual and olfactory cues, especially when it comes to foraging for food. These animals use their vision to find feeding locations and it is hypothesized they use olfactory cues to determine ripeness of fruit.

Scent-marking is a common way of communicating among ruffed lemurs. Females scent-mark by squatting and rubbing their anogenital region back and forth on a substrate. Males sent mark by lowering their body and rubbing their chin, chest, and neck on a surface. Males and females only scent-mark on horizontal surfaces. (Macedonia and Taylor, 1985; Overdorff, et al., 2005; Pereira, et al., 1988; Rushmore, et al., 2012)

Food Habits

Black-and-white ruffed lemurs are mostly frugivorous. Lemurs are 92 percent frugivours, because lemurs are dependent on the high fat and non-structural carbohydrate (NSC) content found in fruit. In seasons when fruit is scarce, the lemurs resort to a folivorous diet, consuming young leaves, flowers, and nectar. The diurnal lemurs spend most of their day foraging for food in mature canopy trees, which produce abundant amounts of fruit. Infant black-and-white ruffed lemurs nurse from their mothers until about 3 or 4 months of age. At this point they are able to forage for their own food. (Schmidt, et al., 2010; Schmidt, et al., 2010)

  • Plant Foods
  • leaves
  • seeds, grains, and nuts
  • fruit
  • nectar

Predation

Lemurs stay high up in the canopies of trees. This could be an evolutionary trait to avoid predators. Lemurs let out warning vocalizations when there is a nearby predator in their territory. One of the sounds is a growl-snort sound that resembles the sound of a dog and the other sound is a pulsed squawking sound. Humans Homo sapiens are a main predator of ruffed lemurs by hunting them for consumption. Raptors are also a main predator of lemurs. Harrier hawks Polyboroides typus are very large and are strong enough to capture and prey on lemurs. (Karpanty and Wright, 2007; Pereira, et al., 1988)

Ecosystem Roles

Black-and-white ruffed lemurs can be infected by the roundworm Baylisascaris procyonis, which causes cerebral nematodiasis. This parasite can cause fetal CNS disease. The paracitic larvae cause severe inflammatory responses in the brain. Lemurs can also be infected by the louse fly, Allobosca crassipes. This ectoparasite only infects lemurs and is flightless as an adult.

Lemurs help their ecosystem by eating fruits and dispersing the seeds when they defecate. (Campbell, et al., 1997; Vaughn, 2010)

  • Ecosystem Impact
  • disperses seeds
Commensal/Parasitic Species

Economic Importance for Humans: Positive

Black-and-white ruffed lemurs are a source of food, though it is illegal to hunt them. They are also in zoos, which is educational and a source of ecotourism. (Andriaholinirina, et al., 2014)

Economic Importance for Humans: Negative

There are no known adverse economic effects of black-and-white ruffed lemurs on humans.

Conservation Status

According to the IUCN Red List, black-and-white ruffed lemurs, and their subspecies white-belted ruffed lemurs Varecia variegata ssp. subcincta, and southern ruffed lemurs Varecia variegata ssp. editorum are critically endangered. In the last 21 years, the population size of ruffed lemurs has declined ≥80%. This is due to habitat destruction such as slash-and-burning of agriculture, logging, and mining. Logging trails that lead deep into the rain forest have made it easier for illegal poachers to access the rain forest. Ruffed lemurs are illegally hunted for consumption.

There are conservation sites and zoos that house ruffed lemurs. Two of the main conservation sites for this species are Betampona Reserve, located northwest of the major port of Tamatave in Madagascar, and Nosy Mangabe, which is an island located off the eastern coast of Madagascar. (Andriaholinirina, et al., 2014)

Contributors

Meg Hale (author), Radford University, Karen Powers (editor), Radford University, April Tingle (editor), Radford University, Emily Clark (editor), Radford University, Cari Mcgregor (editor), Radford University, Jacob Vaught (editor), Radford University, Tanya Dewey (editor), University of Michigan-Ann Arbor.

Glossary

acoustic

uses sound to communicate

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

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

folivore

an animal that mainly eats leaves.

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.

introduced

referring to animal species that have been transported to and established populations in regions outside of their natural range, usually through human action.

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.

nomadic

generally wanders from place to place, usually within a well-defined range.

polyandrous

Referring to a mating system in which a female mates with several males during one breeding season (compare polygynous).

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.

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

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

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