Critically endangered (V. variegata), Endangered (V. rubra)
Life span: 36 years
Total population: 1,000 to 10,000 (V. variegata), 29,000 to 52,000 (V. rubra)
Gestation: 106 days
Height: 43 to 57 cm (M & F)
Weight: 3.6 kg (M), 3.3 kg (F)
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).
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
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).
CURRENT RANGE MAPS (IUCN REDLIST):Varecia rubra
| Varecia variegata
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).
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).
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
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).
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 2014 March 8.