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Ring-tailed lemur
Lemur catta

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SOCIAL ORGANIZATION AND BEHAVIOR

Lemur catta
Photo: Herbert Gustafson

Ring-tailed lemurs live in multi-male/multi-female groups of about 11 to 17 animals but which range in size from six to 35 at Beza Mahafaly Special Reserve and four to 31 at Berenty Private Reserve (Sussman 1991; Hood & Jolly 1995; Sauthers pers. comm.). Ring-tailed lemurs exhibit female philopatry and the core of the social group is the dominant matriline (Jolly 2003). Because females almost always remain in their natal groups while males emigrate into a new group, ring-tailed lemur social groups are centered around a group of related adult females and their offspring. There is usually a single, top-ranking female that initiates the direction of the group and is the focal point of the rest of the group (Jolly 1966; Sauther & Sussman 1993). Often there is more than one matriline in a social group, and pairs of closely related females (mother-daughter and sister-sister) exhibit friendly social interactions including maintaining close spatial proximity and grooming each other frequently while distantly related or unrelated females are more likely to have aggressive encounters (Taylor & Sussman 1985; Sauther & Sussman 1993; Nakamichi & Koyama 1997). There is a dominance hierarchy among females according to matriline and adult females are always dominant over adult males, reinforcing their dominance through agonistic encounters including lunging, chasing, cuffing, grabbing, and biting males. Submissive responses to these aggressive behaviors include jumping away, fleeing, and squealing (Jolly 1966; Taylor & Sussman 1985; Sauther et al. 1999). The hierarchy among ring-tailed lemur females is not linear; daughters do not always assume the rank of their mothers. One explanation for this pattern is that ring-tailed lemur mothers do not support their daughters in agonistic social interactions therefore the daughters do not inherit the rank but fight to achieve their own rank (Nakamichi & Koyama 1997).

There are usually one to three central, or high-ranking, adult males in a ring-tailed lemur social group and several peripheral males. Rank among males is correlated with age (Sussman 1992). The central males are usually in their "prime," estimated to be between six and nine years of age based on dental wear, while the peripheral males are usually recently transferred males, old males, or young adult males that have not yet left their natal group (Sussman 1991; 1992; Gould 1996; Sauther et al. 1999). Males have a separate dominance hierarchy from females and the central males are dominant over the peripheral males, reinforcing the hierarchy through agonistic interactions (Jolly 1966; Sussman 1992). The males are referred to as central and peripheral not just because of their relationships with females, but also their spatial proximity within the group. When a group of ring-tailed lemurs moves from one location to another the highest-ranking females, juveniles, and dominant males lead the procession, while the lower-ranking males lag behind. Once the group has reached its destination and either rests or forages, the low-ranking males continue to remain on the periphery of the group, foraging alone or resting together (Jolly 1966).

Lemur catta
Photo: Roy Fontaine

One of the benefits to being a high-ranking male is increased social interaction with the high-ranking females which may confer benefits such as decreased risk of predation, increased access to food resources, and increased access to reproductively receptive females (Sussman 1992; Gould 1996). Males achieve high-ranking status when they transfer into a group and fight for dominance with the resident males. Young ring-tailed lemur males leave their natal groups in pairs or trios between three and five years of age. The pair will attempt to join a group together, but full assimilation into the group may take many months as group members of both sexes constantly challenge them. Once a young male has successfully joined a group, he remains peripheral and low-ranking (Sussman 1992). Males between the ages of three and four years transfer on average once every 1.4 years while older males, in their prime, settle down and transfer once ever 3.5 years (Sussman 1992). Regardless of how often they transfer, males consistently immigrate into new groups during the six-month period between December and May, and transfers are especially concentrated during the two- to three-week breeding season in April (Sauther 1991; Sussman 1992; Jolly et al. 1993).

When immigration and recruitment causes a ring-tailed lemur social group to get too large, usually greater than 15 to 25 animals with eight to 10 females, the group breaks apart and forms new, smaller groups (Hood & Jolly 1995; Jolly et al. 2002; Gould et al. 2003). In a dramatically seasonal environment with large variation in available resources, large group size can be detrimental to individuals within the group; by splitting into smaller groups, there is less direct competition for resources and the fitness of each individual increases (Hood & Jolly 1995). When groups split, members of the dominant matriline evict members of a subordinate matriline by targeting severe aggression at lower-ranking females and their offspring. Members of the subordinate matriline eventually are pushed out of the group and either start a new group or, very rarely, join a new social group (Sussman 1991; Hood & Jolly 1995). These newly-formed groups have fewer members, which puts them at a disadvantage. Larger groups can displace smaller groups from food patches, have higher protection from predators, and are more likely to dominate in intergroup encounters (Hood & Jolly 1995). When ring-tailed lemur groups encounter each other, a frequent occurrence given their highly overlapping home ranges, the females of each group face each other and confrontations can involve intense staring and glaring, or can escalate to lunging, cuffing, and biting. When these confrontations intensify, they can result in serious injuries or even death (Jolly et al. 1993). At the end of intergroup encounters in the area of home range overlap, both groups retreat to the center of their respective home ranges (Jolly et al. 1993).

REPRODUCTION

Lemur catta
Photo: Roy Fontaine

The reproductive success of ring-tailed lemurs is highly dependent on environmental conditions. In exceptionally good years in the wild, age of maturation is earlier and birthrate and infant survival are much higher compared to years with severe environmental conditions such as drought (Gould et al. 2003; Jolly 2003). In normal years, males and females reach adult size until three years of age, and do not reproduce until between 2.5 to four years of age (Jolly 1966; Gould et al. 2003). Female ring-tailed lemurs at Berenty Private Reserve are more likely to mature and give birth at an earlier age compared with those found at Beza Mahafaly Special Reserve. At Berenty, the lemurs live in a slightly richer environment due to supplemental feeding, the presence of introduced fruit trees, and water provisioning (Gould et al. 2003). Female ring-tailed lemurs in captivity consistently give birth at two years of age (Sussman 1991).

Females are sexually receptive for one to two days each year, and estrus may be as short as six to 24 hours (Van Horn & Resko 1977; Sauther et al. 1999). They exhibit ovarian synchronicity so that all of the adult females in a forest are in estrus at roughly the same time. In the wild, the breeding season lasts between seven and 21 days in May and both males and females have multiple mates (Pereira 1991; Sauther 1991; Sussman 1991). During these few weeks, males approach females to inspect their genitals and attempt mating. Females that are not receptive will act aggressively toward the males, cuffing or chasing them away (Koyama 1988; Sauther 1991). High-ranking males are able to maintain close proximity to females during the breeding period and approach, sit near, and rest or sleep in contact with females throughout the day (Sauther 1991). Estrous females actively approach males for mating by orienting their backsides to a male, lifting their tail, and looking over their shoulder at him. The order of mating reflects the male dominance hierarchy , the highest-ranking, central male is the first male to successfully approach and mate when a female becomes receptive. He is followed by the second-ranking male and then by transfer males or non-troop males. Females reject the mating attempts of related males and sometimes seek out males from other troops, though other males attempt to disrupt these extra-group copulations (Sauther 1991). Aggressive encounters between males increases greatly before and during the breeding season as they fight for access to receptive females (Jolly 1966; Koyama 1988; Sauther 1991).

Lemur catta
Photo: Herbert Gustafson

Because of the harsh environments in which they live, ring-tailed lemurs have high fecundity. Gestation lasts from 135 to 145 days and females in the wild almost always give birth to singletons (Sauther 1991; Koyama et al. 2001; Gould et al. 2003). In captivity the rates of multiple births, including twins and triplets is higher than in the wild (Sussman 1991). Between 75 and 80% of the adult females in the population give birth each year and the average interbirth interval is 1.2 years (Koyama et al. 2001; Jolly et al. 2002; Gould et al. 2003). Having such high rates of reproduction allows ring-tailed lemur populations to rebound after years of high mortality associated with environmental stress. For example, in years of drought, infant mortality can be as high as 80% while normal infant mortality within one year after birth is around 37% (Gould et al. 1999; Koyama et al. 2001; Jolly et al. 2002).

PARENTAL CARE

Ring-tailed lemurs are strict seasonal breeders and give birth to coincide with the end of the dry season and the beginning of the wet season, with the most number of births occurring in September (Koyama et al. 2001). By timing births during periods of high food abundance, some of the physical stress of nursing and weaning an infant may be minimized (Sauther 1998). For the first three weeks of life, the mother is the primary caregiver to her new infant, but she is tolerant of other females in the group, especially siblings of the infant and other mothers (Hosey & Jacques 1994). In the first days of life, infant ring-tailed lemurs cling ventrally to the mother, but by day three, they are able to move actively on her body and can climb onto another grooming female (Jolly 1966). By the age of one month, infant ring-tailed lemurs ride dorsally on their mothers and begin to explore their surroundings independently, spending about 16% of their time off of their mothers. At this age, they do not venture further than .5 m (1.64 ft) from their mothers (Gould 1990). These short stints off of the mother usually involve briefly hopping from her back onto the ground and immediately hopping back onto her. After three weeks, the time spent off of the mother continuously decreases such that by week 16, 85% of the infant's time is spent independently exploring the environment, manipulating objects in the environment, locomotor explanation, and independent foraging (Gould 1990). Because of the timing of births, there are many playmates within the group for infant ring-tailed lemurs. Social play with peers begins around week six and includes such behaviors as chasing, play biting, jumping on, and wrestling with one or more partners (Gould 1990). As the infant matures, it spends less time nursing and more time foraging . Weaning begins at week eight and the mother begins to reject dorsal riding during week 12. By week 16, infant ring-tailed lemurs only nurse about 8% of their total time (Gould 1990).

Other female members of the group, including those with their own new infants, are attracted to newborns and approach mother-infant pairs frequently, attempting to lick or groom infants, especially in the first month of life (Jolly 1966; Gould 1990; Nakamichi & Koyama 2000). These interactions are characterized as affiliative and may serve to increase group cohesiveness. Males within the social group interact with mother-infant pairs much less frequently compared to females, though they are sometimes seen approaching and licking infants. New mothers do not allow strange or newly transferred males to approach them and are extremely aggressive toward any unfamiliar or peripheral males (Nakamichi & Koyama 2000).

Lemur catta
Photo: Herbert Gustafson

In addition to licking and grooming new infants, members of the social group provide varying degrees of infant care for the mother in a behavior called alloparenting (Jolly 1966; Gould 1992). In ring-tailed lemurs, all age- and sex-classes are seen caring for new infants in some capacity. Mothers permit alloparenting by group members because both they and their infants benefit; mothers are allowed time to rest and can forage and travel more efficiently if another ring-tailed lemur is carrying their infant while infants gain valuable social skills, receive additional protection from predators or conspecifics from other groups, and potentially can gain relationships with adult females that will influence their future rank in the group (Gould 1992).

The alloparents also benefit. Adult males gain social access to adult females, potentially securing their position as a mate in the future, nulliparous females gain valuable experience in infant handling and other parental skills, and relatives of the mother and infant increase their inclusive fitness by contributing to the survival of the infant (Gould 1992). Alloparenting is so pervasive that if an infant is orphaned before it is weaned , group members will adopt and provide care for it, including carrying and nursing it (Gould 2000). Incidences of kidnapping have also been seen among ring-tailed lemurs. An adult female, acting as an alloparent , keeps the infant and, with other females, prevents the mother from retrieving it. In some instances, the kidnapper adopts the infant as their own while in other instances, they simply prevent the mother from retrieving it, give the infant no care, and the infant subsequently dies (Koyama et al. 2001).

COMMUNICATION

Olfactory, visual, and vocal communication are important to ring-tailed lemurs. As members of the Order Prosimii, they rely more heavily on olfactory communication than most anthropoid primates, but compared to nocturnal prosimians, visual and vocal signals also play a larger role in social interactions (Gould et al. 1999). Ring-tailed lemurs have 28 distinct call types, 22 of which are used by adults, six of which are particular to infants ( Macedonia 1993). Some predominant vocalizations include affiliative vocalizations such as "moans," contact calls used in conditions of low or moderate arousal, "meows," heard as contact calls in situations of moderate arousal or excitement and which are thought to increase group cohesion, and "wails" which are the highest arousal contact calls and are heard when a group member is separated from the social group. "Howls" are given only by non-infant males and are used to contact and advertise presence of the group to other groups in the area and can be heard between 750 and 1000 m (.466 and .621 mi), "purrs" are heard during grooming and thought to be a sign of contentment, and "chirps" are given to elicit group movement from one location to another (Jolly 1966; Macedonia 1993). Agonistic vocalizations include "yips," given by subordinate animals when approaching or when approached by a dominate individual, "squeals" are given by males during displays to assert status over other males or when soliciting females, and "chutters," given when a dominant individual lunges at a subordinate individual ( Macedonia 1993). Ring-tailed lemurs also have specialized antipredator vocalizations that elicit responses from the rest of the group when they are given. For example, "gulps" are heard when a carnivore, raptor, or rapidly moving human are perceived and are generalized group alert vocalizations, "shrieks" are heard in response to large, low-flying birds, "clicks" are heard in situations of curiosity but wariness, and "yaps" are heard during mobbing of mammalian predators (Sauther 1989; Macedonia 1993).

Lemur catta
Photo: Herbert Gustafson

LISTEN TO VOCALIZATIONS

In addition to vocal communication, ring-tailed lemurs have an array of visual signals including postures that serve to communicate dominance status. A simple visual signal used to intimidate another individual or to start a fight is a "threat stare." The first animal stares at another and the second animal either looks away or will approach and fight. Another visual signal is the "pulled-back lips," done in submission (Jolly 1966; 2003). Jump-fighting is another social interaction involving two animals displaying. This aggressive action involves a ring-tailed lemur standing on its hind legs with arms outspread and jumping or hopping around the other animal involved. This usually occurs on the ground and can escalate, causing serious injury (Jolly 1966). The other typical display seen only among male ring-tailed lemurs is the "stink fight," which involves ritualized posturing (the tails are held over the head and waved back and forth) as well as chemical communication (Jolly 1966).

While males have scent glands on their wrists, on their chests, and in the genital area, females only have anogenital scent glands used for marking (Kappeler 1990; Oda 1999). During a "stink fight," males anoint their tails by rubbing the ends of their tails on the inside of their wrists and on their chests. They then arch their tails over their bodies and wave them at their opponent. The male toward which this is directed either responds with a display of his own, physical aggression, or flees. "Stink fights" can last from 10 minutes to one hour (Jolly 1966). Olfactory communication is also used as a form of intragroup communication as well as intergroup information transfer as ring-tailed lemurs can differentiate between the scents from individual animals. For example, both males and females mark horizontal and vertical substrates at the overlap of their home ranges using their anogenital scent glands (Mertl-Millhollen 1988). In order to mark a vertical substrate such as a tree trunk, they stand on their hands, grasping as high up on the substrate as possible with their feet, and rub their anogenital scent gland along the object (Jolly 1966). Males also leave visual and scent markings using the glands on their inner forearms. This is called "spur marking." They grasp the substrate, usually a small sapling, and drag the thorny nail which overlays the scent gland, cutting into the wood and spreading secretions (Jolly 1966; Mertl-Millhollen 1988). When on the ground, ring-tailed lemurs preferentially mark small saplings and when high in the trees they usually mark small vertical branches (Mertl-Millhollen 1988). Scent markings are left in the area of the home range that overlaps with other groups' home ranges. This area of overlap is where groups are most likely to encounter each other, and since scent marking behavior occurs frequently during intergroup encounters, one function may be to reinforce the boundaries established during confrontations (Mertl-Millhollen 1988). Males "spur mark" more frequently during mating and migration seasons possibly to inform other males of their presence and to deter new males from entering an area to mate with females or to dissuade them from attempting to immigrate into the group. Direct male-male competition is high during the mating season and fights can result in serious injuries, therefore scent marking during the mating season and when new males are likely to enter the social group serves to advertise a male's presence to other nearby males decrease the chances of direct confrontation or dangerous fighting (Gould & Overdorff 2002).

Content last modified: September 21, 2005

Written by Kristina Cawthon Lang. Reviewed by Michelle Sauther.

Cite this page as:
Cawthon Lang KA. 2005 September 21. Primate Factsheets: Ring-tailed lemur (Lemur catta) Behavior . <http://pin.primate.wisc.edu/factsheets/entry/ring-tailed_lemur/behav>. Accessed 2014 December 19.