Ruffed lemur

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).

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SOCIAL ORGANIZATION AND BEHAVIOR

Ruffed lemur social organization is marked by significant variation in both group organization as well as group composition. This has been suggested to be the result of the social system being adaptable to allow for an inability of the species to change its feeding ecology (Vasey 2003). Some of the variation in past studies of ruffed lemur social organization might be attributed to the short-term or seasonal nature of field studies not adequately covering yearlong patterns of social behavior (Vasey 2003). Ruffed lemurs are organized into dispersed social networks which include core groups organized within a core community. Even though the core community lives within a discrete community home range, the entire community is never seen together in the same location at the same time (Morland 1991a; Rigamonti 1993; Vasey 1997a; 2006). The structure of the core group is variable but often consists of two reproductive females in addition to males and subadults, and can range from a pair of individuals up to 9 (Vasey 1997a). Rare interactions between group males are typically agonistic (Morland 1991b). Perhaps the ruffed lemur social structure is best described as multi-male/multi-female fission-fusion social organization (Vasey 1997a; 2006). Such communities are dispersed core social groups that interact with other core groups in the community, however interaction between different core social groups is curtailed during the cold rainy season, the period of gestation and mating (Rigamonti 1993; Vasey 1997a; 2006). During this period of the year, small subgroups are formed and range over small areas within the communal home range (Morland 1991b). Often, these subgroups consist of a male, female, and their offspring or even just an adult male and adult female and might be misconstrued as monogamous (Morland 1991b; White 1989). In captivity, the relationships between mothers and their adult female offspring are not stable throughout the year, with affiliative behavior seen between the two during the birth season but not the mating season (White et al. 1992). The fission-fusion nature of the social organization is manifested on two levels, the yearlong daily changes in subgroups and the seasonal dispersal of core groups into core areas (Vasey 2006). Even though ruffed lemur communities are not cohesive units, the home range is communally defended. In addition, there is some evidence that only females participate in communal home range defense against females from other groups, which includes agonistic behaviors including chasing, scent-marking, calling, and sometimes physical contact with members of neighboring communities (Morland 1991a; 1991b; Vasey 1997a; 2006). Territorial disputes occur most often during the hot months when resources are more readily available and happen at the boundaries between communal home ranges (Morland 1991b; Vasey 2006). During such disputes, males will scent-mark but will not chase, remain uninvolved and are relatively silent (Morland 1991b; Vasey 2006). Group size is very variable, with cohesive groups ranging from a pair to 31 individuals (White 1991; Morland 1991a; Rigamonti 1993; Vasey 1997a; 2003; 2006). In captive and free-ranging ruffed lemurs, females are almost always dominant to males, winning almost all agonistic encounters with them and rarely showing submissive behavior towards a male (Kaufman 1991; Raps & White 1995; Meyer et al. 1999). In the wild, the nature of female group dominance is more ambiguous. While it is possible for females to be dominant, wild groups cannot be described definitively so, as there is some inter-group variation in dominance patterns (Overdorff et al. 2005). In the wild, common aggressive behaviors include attacks, cuffs, grapples, and chases while in captivity the aggressive repertoire includes the stare, charge, chase, lunge, cuff, feint-to-cuff, bipedal hop, pounce on, push down, and bite (Pereira et al. 1988; Overdorff et al. 2005). In the wild, chatter vocalizations are used to signal submission (Overdorff et al. 2005). Captive animals also chatter to show submission in addition to displacement, head turning/eye aversion, cowering/flinching, grimacing, backing away, fleeing, and jumping away (Pereira et al. 1988). In the wild, affiliative behaviors include the greeting behavior performed by females in which their bodies are intertwined. In addition, other wild affiliative behaviors include playing and social grooming. Often, affiliative behaviors are highly seasonal and some are only performed by one sex, such as the male behaviors of the squeal approach and anogenital inspections, which only occur during the mating season (Morland 1991a). In captivity, affinitive social behaviors include group movement, huddling together with bodily contact, greeting by sniffing and social grooming, an activity performed by all group lemurs over 5 months of age (Pereira et al. 1988). In captivity, all group members will play, but subadults participate more often than adults. Play can include wrestling, grappling, chasing, fleeing and solitary play (Pereira et al. 1988).

REPRODUCTION

Wild ruffed lemurs exhibit a seasonal polygamous mating system contrary to initial reports of monogamy in the species (Vasey 2007). In the wild, mating occurs with community members as well as with members from other communities. In addition, a single male or female will often mate with more than one partner in a single mating season and within a community, multiple males and females actively mate (Morland 1993b; Vasey 2007). Near the northern limit of the ruffed lemur range, mating occurs in the cold rainy season, between May and July (Morland 1993b; Vasey 2007). As expected in a species with a specific mating season, births are also seasonal. In the wild on the Masoala peninsula, births occur during the hot dry season during November and are highly synchronized (Vasey 1997a). In captivity, discrete mating and birth seasons are also observed (Brockman et al. 1987). ust prior to and during estrus, both wild and captive females exhibit swelling of the sex skin, peaking around the middle of estrus (Bogart et al. 1977; Brockman et al. 1987; Vasey 2007). Estrous cycles average 14.8 days in captivity (Brockman et al. 1987). Male testicular volume also increases as the mating season nears, peaking prior to or at the time of breeding (Bogart et al. 1977; Foerg 1982; Brockman et al. 1987). Aggression among ruffed lemurs of the same sex increases as well during the mating season (Morland 1993b). Before copulation, the male will perform a display which includes moving toward the female while lowering his head and squealing, roar-shrieking with the female, licking or sniffing the female’s genitals, submissively chattering, mounting and scent-marking. Females also perform displays that include roar-shrieking with the male, hitting, biting, and posturing of her body for mounting (Vasey 2007). The male mounts the female by grasping her hind limbs or a branch with his hind limbs and grasping her torso and thrusting. Aggressive behavior by the female is often directed towards the male who is attempting to mate with her and a mating pair will often copulate a number of times in a bout of mating (Foerg 1982; Morland 1993b). In captivity, gestation averages around 102 days and in the wild, is slightly longer, at 106 days with an inter-birth interval of one year (Bogart et al. 1977; Foerg 1982; Rasmussen 1985; Brockman et al. 1987; Vasey 2007). Reproductive maturity in captivity is reached in both sexes around 18-20 months of age, during the second post-natal breeding season (Foerg 1982; Porton 1989). In the wild, reproductive maturity is reached later, with females only coming into estrus at 3 years old and males showing no signs of reproductive maturity until at least 5 years old (Morland 1991a). Reproductive activity in captivity for females can last into the 23rd year of life (Weigler et al. 1994).

PARENTAL CARE

In both the wild and captivity, ruffed lemur births are usually multiple, averaging around two infants per birth in captivity (Boskoff 1977; Morland 1990; Weigler et al. 1994; Vasey 2007). However, numbers of infants can range from one per birth up to five (Weigler et al. 1994; Greeley 1982; Pollock 1986). Birth weight in captivity ranges from 70 to 140 g (.15 to .31 lb), averaging between 83.0 to 101.7g (.18 to .22 lb) (Boskoff 1977; Brockman et al. 1987). At birth, newborns are covered with fur and their eyes are open (Foerg 1982). Before giving birth, a ruffed lemur mother will construct an arboreal nest within her core area, normally between 10 to 25m (32.8 to 82.0 ft) above the ground, out of branches, leaves and other foliage (Morland 1990; Vasey 1997a; 2007). Such nests are shallow and dish-shaped with only one apparent entry point (Vasey 2007). In captivity, for the first two weeks of life, mothers spend between 70% and 90% of their time in their nests with the newborns (Pereira et al. 1987). In the wild, the infant remains in the nest until about one to three weeks of life, at which point the mother moves them out for the first time and “stashes” them while she performs other activities (Pereira et al. 1987; Morland 1990; Vasey 2007). Infant “stashing” involves placing the infant or infants in concealing foliage in the canopy and leaving them there for up to several hours at a time while she forages elsewhere, sometimes out of earshot (Morland 1990; Vasey 2007; see Vasey 1997a:42-3 for discussion of the term “stashing”). Transport of infants by the mother occurs singly, with the mother grasping the infant’s belly crosswise in her mouth and continues until they are too big to carry, at around 2.5 months old (Morland 1990; Downman 1993; Vasey 2007). While “stashed,” infants will rest, sit still, and not vocalize (Pereira et al. 1987; Vasey 2007). Often, while the mother is away from her offspring, other group members will guard them in addition to emitting alarm calls when sensing danger, responding to the alarm calls of others and providing care for the infants. Males provide a lot of infant care as well; guarding, huddling, grooming and playing with the offspring of up to several different mothers (Vasey 2007). In captivity, females related to a mother have been observed nursing her offspring and close kin have served as foster parents for infants rejected by their mother (Pereira et al. 1987; Kerridge 1999). Indeed, alloparenting is widespread in wild ruffed lemur populations, with all members of the community participating in the raising of offspring (Vasey 2007). In the wild and in captivity, community members take part in “coordinated vigilance displays” in which a community member guarding or near to an infant will alarm call if leaving the infant alone or encountering a predator. Upon hearing the alarm call, other community members will alarm call as well, resulting in the communal transmission of the alarm call throughout the forest and potentially summoning the mother back to her “stashed” offspring (Pereira et al. 1987; Vasey 2007). At one month of age, wild infants will begin climbing and clinging. By two to three months old, the infants will start following their mother and group members for up to 50 to 100m (164.0 to 328.1 ft) and adult mobility and behavior including traveling full-time with adults is attained at 3-4 months old (Morland 1990; Vasey 2007). In captivity, 75-80% of play in the first three months of life is with the mother (Downman 1993). By 10 weeks old, infants begin participating in greeting displays and calling begins at about 16 weeks old (Vasey 2007). Scent marking commences at six months old (Vasey 2007). Also, at around 4 months old in captivity, the infant weighs about 70% of the weight of the adults (Pereira et al. 1987). The wild infant will first sample, but not necessarily ingest, solid foods at around two months old. Wild weaning is estimated to occur at around 4 months of age but some individuals have nursed as late as 7-8 months old (Morland 1990; Vasey 2007). Infant mortality in the wild shows great variance, and can range from a quite high 64%, possibly due to falls from arboreal nests, to zero mortality over a birth season (Morland 1990; Vasey 2007).

COMMUNICATION

Wild ruffed lemurs are extremely noisy and can be heard from about a kilometer away (Morland 1991a). Captive ruffed lemurs have three rough classes of vocalizations, high-, medium- and low-amplitude calls. High-amplitude calls include the contagious “roar/shriek chorus” which probably functions in inter-group spacing and communication as well as social functions and is communally participated in by all wild group members including 3-4 month old infants (Pereira et al. 1988; Morland 1991a; Geissmann & Mutschler 2006). “Roar/shriek choruses” can occur throughout the day but are concentrated during periods of high activity (Morland 1991a; Geissmann & Mutschler 2006). Other calls of this time are “abrupt roars” which might function to alert group members to avian predators, to communicate between group members out of visual contact and also probably to communicate a more general aggressive/defensive response to a disturbance, and “pulsed squawks” which primarily alerts group members to mammalian predators, shows high arousal and is a signal for the group to quickly aggregate (Pereira et al. 1988; Macedonia 1990; Morland 1991a). In the wild, “roar/shriek choruses” and “abrupt roars” are emitted more often during the hot seasons than during the rest of the year (Morland 1991a). Other high-amplitude calls are the “wail,” which functions to bring the group back together or signal all-clear, and the “bray” and “quack,” which serve possible mating functions as a male display (Pereira et al. 1988; Morland 1991a). Loud calls are delivered with the lemur’s body in a “taut” posture (Vasey 2003). Moderate-amplitude calls are the “growl” which alerts the group to a mild disturbance or to the approach of an individual, the “growl-snort” which alerts the group, “chatter” which signals subordinate status, and the “whine” which signals appeasement by males in the mating season or submission (Pereira et al. 1988; Morland 1991a). Low-amplitude calls are the “grunt,” signaling mild annoyance, the “huff,” signaling intense aggravation, especially when avian predators are present, and the “mew,” often exchanged between mother and offspring when separated or during adults during travel (Pereira et al. 1988; Morland 1991a). Other wild vocalizations are the “cough,” which signals aggression between a female and a male during mating and birth seasons, the “grumble” which advertises the presence of a male to another, the “squeak” which infants use to signal distress, and the “squeal” of affiliation between females (Morland 1991a). In the wild and in captivity, scent-marking is also an important means of communication to the ruffed lemur and likely communicates information about sex, location and identity (Morland 1991a). Female ruffed lemurs predominantly use their ano-genital scent glands to mark while males use chest, neck, and mandible glands (Pereira et al. 1988; Vasey 2003). Females mark by squatting and rubbing their ano-genital region over a horizontal surface excepting the ground. Males mark by rubbing their chest, chin, and neck on a horizontal or vertical surface including the ground. Both sexes occasionally mark in the characteristic fashion of the opposite sex (Pereira et al. 1988; Morland 1991a). Scent-marking also plays a role in greeting displays, in which individuals “leapfrog” over one another and scent-mark the back of the other individual (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) Behavior . <http://pin.primate.wisc.edu/factsheets/entry/ruffed_lemur/behav>. Accessed 2020 July 10.

INTERNATIONAL STATUS

For individual primate species conservation status, please search the IUCN Red List. Also search the current scientific literature for primate conservation status (overall as well as for individual species), and visit CITES (Convention on International Trade in Endangered Species of Wild Fauna and Flora).

Conservation information last updated in 2007 follows, for comparison:

CONSERVATION THREATS

Threat: Human-Induced Habitat Loss and Degradation

As with other primates, habitat loss through deforestation is a significant threat to the ruffed lemur. In fact, lemurs are in more grave danger of becoming extinct than most other primates and are most threatened by habitat destruction and hunting (Ganzhorn et al. 1996/1997). Deforestation on Madagascar is undertaken to support subsistence agriculture, cash crops and provide firewood and is especially damaging in river valleys and on the coast (Simons & Lindsay 1987). On the Masoala peninsula, the only habitat of V. rubra, slash and burn agriculture (tavy) is practiced seasonally between October and December and is expanding (Vasey 1996). Also, in some cases, cattle allowed to free-range over former agricultural clearings prevent the re-growth of forest (Evans et al. 1993-1994). However, ruffed lemurs are present in and can survive in very disturbed habitats, albeit probably at far lower densities that in natural habitats (Simons & Lindsay 1987). Selective logging for certain sizes and species of trees affects ruffed lemurs differently than other lemur species because of their reliance on larger trees and the fruits of certain hardwoods. Several species of tree preferred for construction materials are also preferred by ruffed lemurs, so even if a forest is only selectively logged for such materials, the species is still affected. Forest exploitation which does not completely destroy the habitat can also potentially affect ruffed lemur travel routes through the canopy (White et al. 1995).

Threat: Harvesting (hunting/gathering)

Trapping and hunting with traditional weapons of ruffed lemurs occurs as a source of subsistence for local inhabitants (Simons & Lindsay 1987; Evans et al. 1993-1994). Two types of hunting threaten the ruffed lemurs of the Masoala peninsula; using traditional traps across cut swaths of forest (laly) and using firearms (Vasey 1996). The hunting season in this region is between May and September and local attitudes show that while hunting is recognized as illegal, laws are generally not enforced and the local inhabitants are not ashamed about their use of the lemurs (Vasey 1996). Hunting is probably the greatest threat to the lemurs of the Masoala peninsula because while logging and slash and burn agriculture conceivably could be curtailed, hunting would continue (Vasey 1996). Also, even in regions where hunting of lemurs has stopped, hunters pursuing other quarry sometimes adversely affect ruffed lemurs by inadvertently chasing them from favorite food sources. Finally, ruffed lemurs are sometimes also taken alive from their natural habitats as pets or tourist curiosities (Ratsimbazafy 2002).

Threat: Natural Disasters

Tropical cyclones can have a severe impact on ruffed lemurs. In one case, a cyclone destroyed 80% of the forest canopy in a ruffed lemur habitat, severely impacting the large trees the species relies on for food and other aspects of its ecology (Ratsimbazafy 2002). In the ensuing several years, the ruffed lemurs broadened their diet to stave off starvation but remained surprisingly frugivorous. Because of a reduction in body weight resulting from the destruction of food sources, no births were reported for four years among the ruffed lemurs affected by the cyclone (Ratsimbazafy 2002). Thus, tropical storms can represent a significant threat to an already stressed species. It is suggested however that the high reproductive capacity and litter size of the ruffed lemur might be an adaptation to counter such natural occurrences (Ratsimbazafy 2002).

SPECIAL NOTES

Starting in 1997 a captive-bred reintroduction program was started to reinforce a population of black-and-white ruffed lemurs at Betampona natural reserve in eastern Madagascar (Britt et al. 1988). To date, the program has met with some success, including the successful integration of a male individual into a wild group (Britt et al. 2000). In addition, an introduced male and female have both successfully bred with wild individuals (Britt et al. 2003).

LINKS TO MORE ABOUT CONSERVATION

CONSERVATION INFORMATION

• Lemurs in Peril  (National Geographic)
• Monitoring Impacts of Natural Resource Extraction on Lemurs of the Masoala Peninsula, Madagascar
• Links for all species

CONSERVATION NEWS

• Lemurs are key to health of Madagascar’s rainforests  (Mongabay; June 12, 2008)
• Surviving Pioneer Lemurs Celebrate A Decade In The Rain Forest  (ScienceDaily; November 7, 2007)
• Endangered species breed at Welsh farm  (News Wales; May 18, 2004)
• Duke University Lemurs Are Released Into Wilds of Madagascar  (ScienceDaily; November 18, 1997)
• Links for all species

ORGANIZATIONS INVOLVED IN Varecia CONSERVATION

• LVDI International
• Madagascar Fauna Group (MFG)
• Monkeyland Primate Sanctuary
• Rimrock Ranch Wildlife Conservancy, Ltd.
• Wildlife Trust

Content last modified: August 17, 2007

Written by Kurt Gron.

Cite this page as: Gron KJ. 2007 August 17. Primate Factsheets: Ruffed lemur (Varecia) Conservation . <http://pin.primate.wisc.edu/factsheets/entry/ruffed_lemur/cons>. Accessed 2020 July 10.

The following references were used in the writing of this factsheet. To find current references for Varecia, search PrimateLit.

REFERENCES

Balko EA. 1998. A behaviorally plastic response to forest composition and logging disturbance by Varecia variegata variegata in Ranomafana national park, Madagascar. PhD dissertation, State University of New York, Syracuse. Balko EA, Underwood HB. 2005. Effects of forest structure and composition on food availability for Varecia variegata at Ranomafana National Park, Madagascar. Am J Primatol 66(1):45-70. Bogart MH, Cooper RW, Benirschke K. 1977. Reproductive studies of black and ruffed lemurs Lemur macaco macaco and L. variegatus ssp. Int Zoo Yb 17:177-82. Britt A. 2000. Diet and feeding behaviour of the black-and-white ruffed lemur (Varecia variegata variegata) in the Betampona reserve, eastern Madagascar. Folia Primatol 71(3):133-41. Britt A. 1997. Environmental influences on the behavioural ecology of the black-and-white ruffed lemur (Varecia variegata variegata, Kerr 1792). PhD dissertation, University of Liverpool. Britt A, Welch C, Katz AS. 2003. Can small, isolated primate populations be effectively reinforced through the release of individuals from a captive population? Biol Conserv 115(2):319-27. Britt A, Welch C, Katz AS. 1998. First release of captive-bred lemurs into their natural habitat. Lemur News 3:8-11. Britt A, Welch C, Katz A. 2001. The impact of Cryptoprocta ferox on the Varecia v. variegata reinforcement project at Betampona. Lemur News 6:35-7. Britt A, Welch C, Katz A. 2000. Ruffed lemur re-stocking and conservation program update. Lemur News 5:36-8. Brockman DK, Willis MS, Karesh WB. 1987. Management and husbandry of ruffed lemurs, Varecia variegata, at the San Diego zoo. II. Reproduction, pregnancy, parturition, litter size, infant care, and reintroduction of hand-raised infants. Zoo Biol 6(4):349-63. Downman M. 1993. Breeding behaviour in the black and white ruffed lemur Varecia v. variegata at the Cotswold wild life park. Ratel 20(3):87-91. Evans MI, Thompson PM, Wilson A. 1993-1994. A survey of the lemurs of Ambatovaky special reserve, Madagascar. Prim Conserv 14/15:13-21. Foerg R. 1982. Reproductive behavior in Varecia variegata. Folia Primatol 38(1-2):108-121. Ganzhorn JU, Landgrand O, Wright PC, O’Connor S, Rakotosamimanana B, Feistner ATC, Rumpler Y. 1996/1997. The state of lemur conservation in Madagascar. Prim Conserv 17:70-86. Gebo DL. 1987. Locomotor diversity in prosimian primates. Am J Primatol 13(3):271-81. Geissmann T, Mutschler T. 2006. Diurnal distribution of loud calls in sympatric wild indris (Indri indri) and ruffed lemurs (Varecia variegata): implications for call functions. Primates 47(4):393-6. Goodman SM, Ganzhorn JU. 2004. Biogeography of lemurs in the humid forests of Madagascar: the role of elevational distribution and rivers. J Biogeog 31(1):47-55. Greeley ML. 1982. Quintuplets! Zoonooz 55(6):16. Groves C. 2001. Primate taxonomy. Washington DC: Smithsonian Inst Pr. 350 p. Kappeler PM. 1991. Patterns of sexual dimorphism in body weight among prosimian primates. Folia Primatol 57(3):132-46. Karpanty SM. 2006. Direct and indirect impacts of raptor predation on lemurs in southeastern Madagascar. Intl J Primatol 27(1):239-61. Karpanty SM, Grella R. 2001. Lemur responses to diurnal raptor calls in Ranomafana national park, Madagascar. Folia Primatol 72(2):100-3 Kaufman R. 1991. Female dominance in semifree-ranging black-and-white ruffed lemurs, Varecia variegata variegata. Folia Primatol 57(1):39-41. Kerridge FJ. 1999. Part-time fostering by a pair of black and white ruffed lemurs (Varecia variegata variegata). Anim Welf 8:35-42. Lehman SM, Ratsimbazafy J, Rajaonson A, Day S. 2006. Ecological correlates to lemur community structure in southeast Madagascar. Intl J Primatol 27(4):1023-40. Lindsay NBD, Simons HJ. 1986. Notes on Varecia in the northern limits of its range. Dodo, J Wildl Preserv Trusts 23:19-24. Macedonia JM. 1990. What is communicated in the antipredator calls of lemurs: evidence from playback experiments with ringtailed and ruffed lemurs. Ethology 86:177-90. Meyer C, Gallo T, Schultz ST. 1999. Female dominance in captive red ruffed lemurs, Varecia variegata rubra (primates, lemuridae). Folia Primatol 70(6):358-61. Mittermeier RA, Konstant WR, Hawkins F, Louis EE, Langrand O, Ratsimbazafy J, Rasoloarison R, Ganzhorn JU, Rajaobelina S, Tattersall I, Meyers D. 2006. Lemurs of Madagascar. Washington, D.C.:Conserv Intl. 520 p. Mittermeier RA, Konstant WR, Nicoll ME, Langrand O. 1992. Lemurs of Madagascar: an action plan for their conservation 1993-1999. IUCN: Gland, Switzerland. 58p. Morland HS. 1990. Parental behavior and infant development in ruffed lemurs (Varecia variegata) in a northeast Madagascar rain forest. Am J Primatol 20(4):253-65. Morland HS. 1991b. Preliminary report on the social organization of ruffed lemurs (Varecia variegata variegata) in a northeast Madagascar rain forest. Folia Primatol 56(3):157-61. Morland HS. 1993b. Reproductive activity of ruffed lemurs (Varecia variegata variegata) in a Madagascar rain forest. Am J Phys Anth 91:71-82. Morland HS. 1993a. Seasonal behavioral variation and its relationship to thermoregulation in ruffed lemurs (Varecia variegata variegata). In: Kappeler PM, Ganzhorn JU, editors. Lemur social systems and their ecological basis. New York:Plenum Pr. p 193-203.

IMAGES

	Varecia rubra Photo: Bertrand L. Deputte
	Varecia rubra Photo: Pavel Vlcek
	Varecia rubra Photo: Primates in Art & Illustration Collection
	Varecia rubra Photo: Richard Frazier
	Varecia rubra Photo: Rick Murphy
	Varecia variegata Photo: Jennifer Simonson
	Varecia variegata Photo: Jennifer Simonson
	Varecia variegata Photo: Kevin Schafer
	Varecia variegata Photo: Kevin Schafer
	Varecia variegata Photo: Kevin Schafer
	Varecia variegata Photo: Kevin Schafer
	Varecia variegata Photo: M.D. Stuart
	Varecia variegata Photo: Pavel Vlcek
	Varecia variegata Photo: Pavel Vlcek
	Varecia variegata Photo: Pavel Vlcek
	Varecia variegata Photo: Pavel Vlcek
	Varecia variegata Photo: T. Keith-Lucas
	Varecia variegata Photo: T. Keith-Lucas
	Varecia variegata Photo: T. Keith-Lucas