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The Socioecology of the Ringtailed Lemur: Thirty-Five Years of Research

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The Socioecology of the Ringtailed Lemur: Thirty-Five Years of Research 120 Evolutionary Anthropology ARTICLES The Socioecology of the Ringtailed Lemur: Thirty-Five Years of Research MICHELLE L. SAUTHER, ROBERT W. SUSSMAN, AND LISA GOULD Linnaeus’1 original scientific description of Lemur catta, the ringtailed lemur, was ters take place between less closely based on a living animal brought to England in 1749.1 Although there were many related individuals.18,20,21 However, un- brief descriptions of wild ringtailed lemurs,2–5 it was not until Jolly6 wrote her now like cercopithecine monkeys, ringtail classic book, Lemur Behavior, that we had our first detailed description of the natural lemurs rarely form third-party alli- history of these beautiful animals (Fig. 1). Since then, long-term field studies, mainly ances during agonistic encounters.20,21 from two study sites in Madagascar, Berenty and Beza Mahafaly (Fig. 2), as well as Furthermore, reconciliation after con- studies on forest-living groups in captivity at the Duke University Primate Center in flict, which is common among anthro- Durham, North Carolina, have greatly expanded our knowledge of the ecology and poid primates, has not been noted in behavior of this species (Table 1, Box 1). Thirty-five years of research on this species this species, indicating that this is not at these various sites indicates that Lemur catta is proving to be every bit as complex a necessary prerequisite for perma- in its behavior as are many anthropoid primates. This very complexity has been nent social group life in primates.22 reflected in the current controversies and questions concerning the ecology and In free-ranging populations, if a behavior of this species. group becomes too large fission may occur, with one matriline evicting another.8,14,17,18,23 Dividing along mat- SOCIAL ORGANIZATION Groups are centered around a core of rilineal lines is also common among adult females and their offspring.6 As female-bonded cercopithecine mon- Like many cercopithecine monkeys, keys.24–26 One important mechanism Lemur catta lives in multi-male female in Old World monkeys, ringtailed le- for facilitating such fission among ring- resident groups that usually contain mur groups characteristically contain tailed lemurs is episodic targeting ag- 10 to 20 animals.6–8,10 Sex ratios are more than one matriline, with some gression.27 When groups reach a criti- approximately 1:17,8,10,11 and sexual di- matrilines being dominant over oth- cal size, females repeatedly attack and morphism is slight.12 Females usually ers.17–19 More friendly social interac- chase individuals of another matri- remain in their natal group, (but see tions occur between close relatives line, sometimes resulting in the evic- below) whereas males migrate.13–16 and more serious aggressive encoun- tion of the latter.27,28 Although there are no documented cases of individual female transfer, there have been rare Michelle Sauther is assistant professor at the University of Colorado, Boulder. She received her cases of mother and adult daughter Ph.D. from Washington University in 1992. Her research focuses on the socioecology of pairs joining other groups.8,15 At Beza nonhuman primates, with special emphasis on how individual life-history factors contribute to variation in feeding and reproductive strategies. She has studied the ringtailed lemurs of Beza Mahafaly, such a pair entered an ex- Mahafaly since 1987. Her most recent work on this population involves a joint effort to help tant group that contained only four develop a genetic profile to address issues of paternity and conservation for ringtailed lemurs. adult females and became the two She has also done research in Guyana, Panama, Kenya, and Costa Rica. Sauther recently chaired a symposium on the feeding behavior of the Malagasy prosimians at the meetings of top-ranking females. This is not al- the American Society of Physical Anthropologists in 1996. [email protected]. ways the case, however. In another Robert W. Sussman is professor in the department of Anthropology at Washington University, case, the attempt of a mother-daugh- St. Louis. He also is Editor-in-Chief of the American Anthropologist. He received his Ph.D. from ter pair to join a group was repulsed. Duke University and has been conducting research on the lemurs of Madagascar since 1969. He also has done research on primates and conservation in Costa Rica, Panama, Guyana, and They eventually joined a different, Mauritius. Besides primate behavior, ecology and conservation, he is interested in the evolution smaller group that contained only two of primate and human behavior. [email protected] resident females.15 Lisa Gould is assistant professor at the University of Victoria, Victoria B.C. Her research interest In all groups studied at Beza Maha- focuses on ringtailed lemur socioecology, including field studies of infant development, the faly, the top-ranking male has been of structure of male social relationships, and sex differences in vigilance behavior. One goal of 8 these projects is to examine the effect of female dominance on the social system of this prime adult age. Peripheral males are species. Most recently, she has been examining the effect of a severe drought on the not a cohesive subgroup, but include population of ringtailed lemurs at the Beza-Mahafaly Reserve in southwestern Madagascar. lower-ranking males and males at- [email protected] tempting to immigrate into the group. Male migration is regular, with young Key words: Lemur catta; ringtailed lemur; socioecology; reproduction males first migrating from their natal ARTICLES Evolutionary Anthropology 121 Figure 2. Formal field studies of Lemur catta Figure 1. Ringtailed lemurs live in large social groups in the dry and very seasonal forests of began in the 1960s at various sites in Mada- southern Madgascar. gascar. Information from long-term studies at Berenty and Beza Mahafaly are expanding our knowledge of the socioecology of this species. group when they are between 3 and 5 tionships are not invariably linear and years of age.14 On average, mature can change quite abruptly.20,29,33 ranks are actively maintained by fe- males migrate every three-and-a-half There is also a developmental com- male agonistic intervention and daugh- years, with approximately 30% migrat- ponent in the establishment of domi- ters acquire ranks just below those of ing each year.14 Males form partner- nance relationships: Adolescent and their mothers through affiliative inter- ships while transferring, although young adult females may rank above actions and alliances among female these partnerships do not necessarily some adult females.20,32 Perhaps be- relatives.34 It thus appears that al- continue once group membership is cause alliances are rare and mothers though female ringtailed lemurs have attained.13,14,29,30 At Berenty, the major- dominance relationships, they do not ity of male migrations occur at the end maintain dominance hierarchies in the of the dry season and overlap, in part, manner of multi-male groups of cerco- with the birth season.13 In all cases of It thus appears that pithecine primates.32 male transfer, the males initially hold although female Unlike the males of many group- low-ranking and peripheral positions living anthropoid primates, male ring- upon entering a new group and are ringtailed lemurs have tailed lemurs appear to lack a consis- challenged by other group members of dominance tent linear hierarchy and rank reversals both sexes for many months.8,19,29,30 occur.16,18,22,29 This may relate to a lack A single top-ranking female appears relationships, they do not of the stable alliances and coalitions to be the focal point for the rest of the maintain dominance between migrating males that are group. She often initiates the direction found in some macaque and baboon of group progressions, although other hierarchies in the species, but have yet to be observed in individuals, including low-ranking manner of multi-male ringtailed lemurs.32 In most studies, a adult males, may lead the actual single, dominant male could be clearly 6,19 groups of cercopithecine travel. This female is also respon- identified and, at Beza Mahafaly, three sible for the greatest number of aggres- primates. males were able to maintain their domi- sive encounters overall, with the major- nance for six years, a pattern more simi- ity of these involving access to food.31 lar to that of male rhesus macaques than Females show strong, differentiated that of male baboons.8,35 Male ringtailed relationships within matrilines, as seldom support their daughters agonis- lemurs also develop preferred partner- measured by grooming and proxim- tically, female offspring do not invari- ships with one another, although these ity.18–20 Mothers are always dominant ably inherit their mother’s rank.20,32 usually do not persist through an entire over daughters.19,32,33 While female This is different from the relatively year.29,30 (However, see Nakamichi and dominance relationships in ringtailed stable linear dominance hierarchies Koyama.20) lemurs can be linear, transitive, and found among group-living baboons Intergroup encounters are common stable over long periods,19 these rela- and macaques. Among these species, at both field research sites.6,13,19,36–39 122 Evolutionary Anthropology ARTICLES TABLE 1. Main Study Sites of the individuals feeding at the same time 30% of their time feeding on the Ringtailed Lemur, Lemur catta when there is ample food. If a resource ground.10,19 Reproductive state affects (Studies of at Least One Year) is limited—for example, if there is feeding behavior. At Beza Mahafaly, Sites and Dates Main Researchers only a single drinking site—individu- lactating females focus more on low- als may queue up for access to that cost, predictable, high-protein plant Berenty, 1963–pre- A.
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