Ruffed lemur
Varecia

TAXONOMY

Suborder: Strepsirrhini
Infraorder: Lemuriformes
Superfamily: Lemuroidea
Family: Lemuridae
Genus: Varecia
Species: V. rubra, V. variegata
Subspecies: V. v. editorum, V. v. subcincta, V. v. variegata

Other names: ruffed lemur; lémur vari, maki vari (French); lémur de collar, lémur de gola, lémur de gorguera (Spanish); kraglemur, pandalemur, röd vari, svartvit vari, vari (Swedish); V. variegata: black-and-white ruffed lemur; varikandana, varikandra (Malagasy); V. rubra: red-ruffed lemur; varinena, varikamena (Malagasy).

MORPHOLOGY

V. variegata and its subspecies are exclusively black-and-white while V. rubra is predominantly chestnut-red with some black and white as well (Mittermeier et al. 2006). However there is significant pelage variation within the ruffed lemurs and there are intermediates in color between the two species (Mittermeier et al. 2006). In addition, a very abstract pattern of more white and less black in the coloration the further south the population occurs is seen, although only generally (Vasey & Tattersall 2002). The black-and-white ruffed lemur (V. v. variegata) abdomen, tail, hands and feet, inner limbs, forehead, face and crown are black. Pelage is white on the sides, back, hind limbs and on the hindquarters (Mittermeier et al. 2006). V. v. editorum has very similar coloration, except the anterior part of the back which is all black while in V. v. variegata this part of the body has a white band (Mittermeier et al. 2006). V. v. subcincta has black dorsal surfaces with a white "belt" or band around the middle of the body (Groves 2001; Mittermeier et al. 2006). The rest of its body is black except for the outer limbs and base of the tail which is white (Mittermeier et al. 2006). In all species of V. variegata the white fur is sometimes slightly yellowish or even brownish and the black can be slightly grey or brownish (Groves 2001; Vasesy & Tattersall 2002).

Varecia variegata
Photo: Kevin Schafer

The red-ruffed lemur (V. rubra) is immediately very different looking from all varieties of V. variegata, as most of the dorsal surfaces are deep reddish or chestnut-red in color excepting the white nape of the neck (Groves 2001). V. rubra undersides, extremities, face, top of the head and tail are black (Groves 2001). In all species of ruffed lemur, the characteristically long face is black or mostly black contrasting with long, furry "ruffs" of hair on the ears which are white in V. variegata and deep reddish or chestnut-red in V. rubra. Pelage is typically quite fluffy (Vasey 2003; Mittermeier et al. 2006). No sexual dimorphism is seen between males and females nor are differences in coloration evident between the sexes (Kappeler 1991; Vasey 2003). Females possess three pairs of mammary glands (Tattersall 1982).

The ruffed lemur is the largest living member of the Lemuridae family (Vasey 2003; Mittermeier et al. 2006). Body length ranges from 43 to 57 cm (16.9 to 22.4 in) and the tail is longer than the body, averaging around 60 cm long (23.6 in) (Vasey 2003). Wild female ruffed lemurs range from 2.6 to 4.0 kg (5.7 to 8.8 lb), weighing an average of 3.3 kg (7.3 lb), while males range between 2.8 and 4.1 kg (6.2 and 9.0 lb), averaging 3.6 kg (7.9 lb) (Vasey 2003). With both sexes combined, wild ruffed lemurs average 3.6 kg (7.9 lb) and captive ruffed lemurs average 3.5 kg (7.7 lb) (K. E. Glander & E. Balko, unpublished data cited in Terranova & Coffman 1997; Terranova & Coffman 1997). In captivity, obesity is sometimes a problem with weights in European zoos reaching averages as high as 4.3 kg (9.5 lb) (Schwitzer & Kaumanns 2001).

The ruffed lemur is both terrestrially and arboreally quadrupedal (Pereira et al. 1988; Mittermeier et al. 2006). While walking on the ground, the head is typically pointed towards the ground and the tail is held high. Running is performed in a bounding movement (Pereira et al. 1988). When transitioning between trees, leaping is the most common form of movement and the lemur will look over its shoulder while clinging, push off, and twist in the air so that the ventral surface of the body makes contact with the new tree (Pereira et al. 1988). The most common types of movement are quadrupedalism, leaping, clinging, and suspension followed by low occurrences of bridging, bimanual movement, and bipedalism. Compared to other lemurs, suspensory movement is more common in ruffed lemurs and the most common form of movement is above-branch quadrupedal locomotion (Gebo 1987). Resting postures include hunched sitting and upright postures, as well as prone resting on its belly or sunbathing on its back with limbs outstretched (Morland 1993a).

Individuals of both species of ruffed lemur have lived as long as an estimated 36 years (Weigl 2005).

RANGE

In general, ruffed lemurs are found in the eastern rainforests of Madagascar (Mittermeier et al. 2006). Of the two species, the range of the black-and-white ruffed lemur (V. variegata) is far larger, extending in a line down the eastern Malagasay coast from a southern limit around the Mananara River near Vangaindrano to a northern limit somewhat north and west of Maroantsetra, on the Bay of Antongil (Tattersall 1982; Vasey & Tattersall 2002). The northern limit of V. variegata is near the Antainambalana River, ending in a possible zone of hybridization between V. variegata and V. rubra, extending between the Vohimara and Mahalevona Rivers north of the Bay of Antongil (Vasey & Tattersall 2002). The primary range of the red ruffed lemur is the Masoala peninsula east of the Bay of Antongil, extending west to the Antainambalana River where their range meets up with the range of V. variegata (Tattersall 1982; Vasey 2003). To the north of the Masoala peninsula, the range of the red ruffed lemur may extend only to the Cap Est (Tattersall 1982). The northern barrier limiting the distribution of V. rubra is the Lokoho River (Goodman & Ganzhorn 2004). The total wild population of red-ruffed lemurs (V. rubra) is estimated at between 29,000 and 52,000 individuals (Vasey 1997b). The total population of black-and-white ruffed lemurs is probably between 1,000 and 10,000 (Mittermeier et al. 1992).

HABITAT

Ruffed lemurs are limited to the seasonal eastern rainforests of Madagascar and are not particularly flexible when it comes to habitat choice (Vasey 2000a; 2005). Primarily arboreal, ruffed lemurs prefer the crowns of large and taller trees and spend the majority of their time from 15 to 25 m (49.2 to 82.0 ft) above the forest floor (Lindsay & Simons 1986; Vasey 2000a; 2003). Crowns of trees are a primary habitat through all seasons (Vasey 2004). Throughout the ranges of both species of ruffed lemur, seasonal availability of resources is similar. Fruit, flowers, and young leaves are available in the hot seasons with more young leaves and flowers at the end of the cold wet seasons (Vasey 2002; 2006). In the hot rainy season, major branches of trees are favored by both sexes while females prefer lianas in the crowns of trees, a pattern followed by females in the hot dry season as well (Vasey 2002). In the cold season, the use of the crowns of trees is ubiquitous (Vasey 2002).

Varecia rubra
Photo: Rick Murphy

On the Masoala peninsula, the main habitat of the red-ruffed lemur, there are four seasons, hot and rainy (January-March), transitional cold (April-May), cold rainy (June-August), and hot and dry (October-December) (Vasey 2006). On the peninsula, in the Masoala National Park, average annual rainfall is around 5110.26mm (201.2 in) with temperatures ranging between highs of 22.5 to 31.6 °C (72.5 to 88.9 °F) and lows of 19 to 23.5 °C (66.2 to 74.3 °F) (Vasey 2006). This region experiences heavy rainfall between January and August with over 550mm (21.7in) per month during this period (Vasey 2002). As expected, there is some variability between study sites, with the seasons on Nosy Mangabe, near the Masoala peninsula in the Bay of Antongil, being divided into three seasons; cool-wet (May-August), dry season (September-November) and warm-wet (December-April) (Morland 1993). Rainfall on Nosy Mangabe averages 3709mm (146. 0 in) annually over an average of 250 days of rain with the lowest monthly temperatures averaging 21.0 °C (69.8 °F) in July and August and the highest around 26.3 °C (79.3 °F) in February (Morland 1993). Near the southern end of the black-and-white ruffed lemur distribution in the Manombo Classified southeastern lowland rainforest, the habitat is characterized by high rainfall throughout the year with a peak in January through March (Ratsimbazafy 2006). Ruffed lemurs have been sighted at altitudes up to 1353 m (4439 ft) (Lehman et al. 2006).

ECOLOGY

Ruffed lemurs are predominantly frugivorous, in fact the most frugivorous of the lemurs, and will often feed on over 80 and up to 132 plant species (White 1991; Morland 1993; Rigamonti 1993; Vasey 2000b; 2003; 2004; Ratsimbazafy 2006). Feeding time is spent between 74-90% on fruit, 4-21% on nectar, 3-6% on flowers, 1% on mature leaves, and 3-6% on young leaves (Vasey 2003). However, while there is often a large diversity of consumed plant species, typically only several are consumed or utilized as a large proportion of the diet (Rigamonti 1993; Vasey 2000b; Balko & Underwood 2005). Between seasons, the diet exhibits no differences save for females eating more young leaves than males and fewer flowers during the hot dry season (Vasey 2002). When it comes to choice of fruit tree, the availability and accessibility of edible fruit is more important than the species of tree itself (Balko & Underwood 2005). During the hot seasons, the size of food patches is larger than in the cold rainy season, but not as large as during the transitional cold season (Vasey 2002). In addition, significantly less time is spent active in the cool-wet seasons than in the rest of the year (Morland 1993). While pregnant and while lactating, female ruffed lemurs will eat more flowers and young leaves late in the day. These high protein foods offset the high energy cost of reproduction (Vasey 2004). While diets vary at different location, common food plants are Canarium (Burseraceae), Cryptocarya, Ocotea, and Ravensara (Lauraceae), Ficus (Moraceae), Eugenia/Syzygium (Myrtaceae), and Grewia (Tiliaceae) (Vasey 2003). At some locations, ruffed lemurs will descend from trees to consume soil and also occasionally eat fungi (Britt 2000; White 1991).

Annually, ruffed lemurs spend an average of 28% of their time feeding, 53% resting, and 19% traveling with females resting less and feeding more than males. They are least active during the cold seasons. Over the course of the day there is no significant variation in activity budget although there is slightly more rest at midday (Vasey 2005). Time spent feeding over the course of the day is similar throughout the year and ruffed lemurs are found mainly in the crowns of trees all day long (Vasey 2004). In order of preference, ruffed lemurs spend the most time from 15 to 20m (49.2 to 65.6 ft), followed by 20 to 25m (65.6 to 82.0 ft) and finally 10 to 15m (32.8 to 49.2 ft) and they will move lower in the canopy to regulate their body temperature during the hotter seasons (Vasey 2004). In the cool months, 2% of resting time is spent sunbathing, while in the warm months its occurrence is reduced (Morland 1993a).

Varecia variegata
Photo: Jennifer Simonson

Home range can vary widely, ranging from .162 km² (.06 mi²) to a quite large 1.97 km² (.76 mi²) home range with values ranging significantly between (White 1991; Morland 1991a cited in Vasey 2003; Rigamonti 1993; Britt 1997 cited in Vasey 2003; Vasey 1997a; Balko 1998 cited in Vasey 2003; Vasey 2003; 2006). Average female annual home ranges are typically larger than those of the males although during mating and gestation, they become smaller than the home ranges of males (Vasey 1997a; 2006). In other words, female ranging varies by season while male ranging does not (Vasey 1997a). Average daily path is variable also, and varies from 436m to 2250m (1430.4 to 7381.9 ft), in one study averaging 1129m (3704.1 ft) per day (White 1991; Rigamonti 1993).

Raptors are predators of ruffed lemurs, but likely at a low rate, with evidence of predation on ruffed lemurs by Henst's Goshawk (Accipiter henstii) (Karpanty & Grella 2001; Karpanty 2006). The fossa (Cryptoprocta ferox) presents a potential but rare threat to ruffed lemurs, but has not been confirmed to prey upon wild individuals (White 1991; Britt et al. 2001). However, this is likely due to the wild ruffed lemur remaining high in the forest strata the majority of the time. Released captive-bred ruffed lemurs have been preyed upon by fossa as a result of their inexperience with predators (Britt et al. 2001). Because ruffed lemurs nest with young, this makes them potentially susceptible to predation from carnivorous mammals, such as the ring-tailed mongoose (Galidia elegans) and the brown-tailed mongoose (Salanoia concolor) (Vasey 1997a).

Ruffed lemurs can found living in sympatry with a number of other primate species including the greater dwarf lemur (Cheirogaleus major), grey gentle lemur (Hapalemur griseus griseus), sportive lemur (Lepilemur mustelinus), diademed sifaka (Propithecus diadema), brown lemur (Eulemur fulvus), red-bellied lemur (Eulemur rubriventer), eastern avahi (Avahi laniger), indri (Indri indri), rufous mouse lemur (Microcebus rufus) and probably the aye-aye (Daubentonia madagascariensis) (Vasey 1997a; Lehman et al. 2006). Ruffed lemurs probably demonstrate feeding dominance over red-bellied lemurs (Eulemur rubriventer) (Evans et al. 1993-1994). In addition, sympatric grey bamboo lemurs (Hapalemur griseus griseus) avoid ruffed lemurs when they encounter them (Evans et al. 1993-1994). Infant ruffed lemurs have even been observed to play with white-fronted lemurs (E. fulvus albifrons) (Vasey 2007).

Content last modified: August 17, 2007

Written by Kurt Gron.

Cite this page as:
Gron KJ. 2007 August 17. Primate Factsheets: Ruffed lemur (Varecia) Taxonomy, Morphology, & Ecology . <http://pin.primate.wisc.edu/factsheets/entry/ruffed_lemur/taxon>. Accessed 2013 May 23.