Discrimination by male ring-tailed Lemurs (Lemur catta) between the scent marks of male and those of female conspecifics

Olfaction plays an important role in the social communication of all prosimians. (The experiment reported in this paper forms part of an intensive chemobehavioral study of olfaction in Lemur catta (ring-tailed lemur) being carried out in this laboratory.) Five male Lemur cattawere tested on their behavioral responses to paired scent stimuli. Responses measured were (1) total investigation time, (2) arm-marking, (3) ABO/BO rubbing, and (4) flehmen. Males showed a strong discrimination between the scent stimuli,giving higher levels of response to female scent on measures 1, 3, and 4. This response suggests an olfactory-related preference by males for female scent under controlled conditions. This preference may be a consequence of the females’ dominance over males and the brevity of estrus in L. catta,both of which would favor such choice behavior.     

Social Relationships Among Ring-Tailed Lemurs (Lemur catta) in Two Free-Ranging Troops at Berenty Reserve, Madagascar

We observed two free-ranging troops of ring-tailed lemurs at the Berenty Reserve, Madagascar. Kinship affinities in these troops are known only for mothers and their offspring 4 years of age. We attempted to quantify social relationships. Almost all agonistic interactions were dyadic, and triadic agonistic interactions, such as alliances, were very rare. Dominance hierarchies in both sexes in the two troops were not linear. As in cercopithecine monkeys, mothers were dominant over their adult daughters. However, the daughters were not ranked immediately below their mothers. Close proximity and social grooming occurred more frequently between closely related females, such as mother–daughter and sister–sister dyads, than between unrelated females. Frequent-proximity relations also occurred between adult males that had emigrated from another troop and entered the present troop together, even though they did not rank closely to one another. Subordinates were likely to groom and to greet dominants more frequently than vice versa. During group encounters, particular females were involved in agonistic interactions with animals of other troops, regardless of dominance rank. Adult males, regardless of their dominance rank, but not adult females, constantly tried to drive solitary males away.     

A preliminary study of kinship and social organization in a semi-free-ranging group ofLemur catta

From long-term studies of a number of anthropoid species, many investigators have shown that kinship affinities affect social relationships. Factors such as proximity, social grooming, dominance rank, and mating patterns have been shown to be related to kinship. In this paper, we report the results of a preliminary study of the social organization of a group of prosimians (Lemur catta) in which individuals were identified and kinship affinities were known. We found that close matrilineal kin preferred to groom one another and to remain in close proximity more than did nonkin and distantly related animals. Furthermore, no copulations were observed within matrilines. These results are similar to those found in a number of species of anthropoids. This research was conducted on a semi-free-ranging group at the Duke University Primate Facility, Durham, North Carolina.     

Development of Olfactory Behavior in Captive Ring-Tailed Lemurs (Lemur catta)

We conducted a long-term research project (1996–1999) on developmental aspects of olfactory behavior in ring-tailed lemurs to document the ontogenetic sequence of olfactory behavior, including the possible presence of sexual differences, and the maturation of scent-marking. The subjects were a group of 18 lemurs housed in the Pistoia Zoo (Tuscany, Italy), which we observed during 1,735 h via all-occurrences sampling, scan-animal sampling and ad libitum sampling methods. We determined the time sequence of olfactory exploration and of scent-marking patterns, and investigated sexual differences in timing and frequency. We also followed the development of scent-marking through the juvenile and adolescent phases taking into account the two aspects of motor control and of the acquisition of social competence. On the whole, we found that olfactory investigation appears and matures earlier than scent-marking. Moreover, olfactory investigation of conspecifics appeared later than substrate exploration, and seemed to follow a sequence of increasing level of acceptance by the receiver. Social play is very important for the maturation of the gestural component of scent-marking. The olfactory behavioral pattern appeared to mature during the juvenile and adolescent phases. Although sexual maturation had obvious influence on the development of olfactory behavior, the onset of scent-marking patterns was only partially parallel to sexual maturation.     

Evidence that displacement activities facilitate behavioural transitions in ring-tailed lemurs

Displacement activities are behavioural patterns defined by their apparent irrelevance to an animal's ongoing actions. Despite being identified in diverse taxa, their function remains poorly understood. One hypothesis posits that displacement activities facilitate transitions between different behaviours by mediating changes in animals' motivational state. Under this hypothesis, it is predicted that displacement activities will occur more frequently around changes in behaviour than at other times, and also that rates of displacement activities will be higher before than after such behavioural transitions. We tested these two predictions in wild ring-tailed lemurs (Lemur catta). During focal observations, animals' behavioural state was continuously recorded, as were all occurrences of self-scratching, a common displacement activity in this species. Self-scratching rates were found to be significantly elevated both before and after behavioural transitions. Furthermore, self-scratching rates were significantly higher before behavioural transitions occurred than after. These results, therefore, provide support for the hypothesis that displacement activities facilitate behavioural transitions in L. catta.     

Gait mechanics of lemurid primates on terrestrial and arboreal substrates

Aspects of gait mechanics of two lemurid species were explored experimentally. Substrate reaction forces were recorded for three animals each of L. catta and E. fulvus walking and running at voluntary speeds either on a wooden runway with an integrated force platform or on elevated pole supports with a section attached to the force platform. The average height of the back over these substrates and fluctuations in this height were evaluated using video-analysis. Animals preferred walking gaits and lower speeds on the poles, and gallops and higher speeds on the ground. At overlapping speeds, few adjustments to substrate types were identified. Hind limb peak forces are usually lower on the poles than on the ground, and the caudal back is closer to the substrate. This suggests that greater hind limb flexion and reduced limb stiffness occurred on the poles. The support phases for both limbs at higher speeds are slightly elongated on the poles. Forelimb peak forces are not lower, and the trajectory of the caudal back does not follow a smoother path, i.e., not all elements of a compliant gait are present on the simulated arboreal substrates. The horizontal, rigid poles, offered as substitutes for branchlike supports in the natural habitat, may not pose enough of a challenge to require more substantial gait adjustments. Across substrates, forelimb peak forces are generally lower than hind limb peak forces. The interlimb force distribution is similar to that of most other primates with more even limb lengths. Walking gaits present a greater divergence in fore- and hind limb forces than galloping gaits, which are associated with higher forces. The more arboreal E. fulvus has higher forelimb forces than the more terrestrial L. catta, unlike some anthropoid species in which the arborealists have lower forelimb forces than the terrestrialists. As in other primate and nonprimate quadrupeds, the major propulsive thrust comes from the hind limbs in both lemurs. While our data confirm certain aspects of primate gait mechanics (e.g., generally higher hind limb forces), they do not fully support the notion of greater limb compliance. Neither a compliant forelimb on branchlike supports, nor a negative correlation of forelimb force magnitudes with degree of arboreality were observed. Increasing forelimb-to-hind-limb-force-ratios with increasing speed and force magnitudes are also not expected under this paradigm.     

Foraging Success,Agonism, and Predator Alarms: Behavioral Predictors of Cortisol in <Emphasis Type="Italic">Lemur catta</Emphasis>

Primate social grouping is understood as an adaptive strategy for mitigating environmental selection pressures, but the relative importance of various pressures may vary. Physiological measures of well-being can show their short-term impacts and suggest their relative importance and capacity to provide ultimate or proximate control of group size. I examined correlations between pressures commonly proposed as causes of social grouping (foraging success, intergroup and intragroup agonism, and predation risk) and individual levels of fecal cortisol, a hormonal stress measure, in a free-ranging population of Lemur catta. I collected behavioral data on 45 female Lemur catta at Berenty Reserve, Madagascar, over 3 seasons (August 1999-July 2000) and determined individual cortisol levels from 474 fecal samples. Neither predator alarm rates nor intragroup agonism rates correlated with cortisol levels in any season. However, females with low daily food intake and high rates of escalated intergroup defense exhibited higher cortisol levels. The data suggest that acquisition and defense of food resources are principal challenges in Lemur catta, and may be important factors determining social grouping and other behavioral or life history adaptations.     

Aggression and Reconciliation in Two Captive Groups of <Emphasis Type="Italic">Lemur catta</Emphasis>

Kappeler excluded the presence of reconciliation in a free-ranging group of Lemur catta, but a recent study on the same group indicated reconciliation, though with a very long PC duration. We collected data on 2 captive groups of ring-tailed lemurs at the Pistoia Zoo (Italy) in order to shed light on conflict resolution in the species. We investigated the influence of seasonality and of a targeted aggression episode on the occurrence of reconciliation. We collected 164 PC-MC pairs for the A group and 141 for the B group. We performed all analyses at the dyadic level via randomization procedures. Despite the targeting episode, we found no difference in the levels of aggression between the 2 groups. In contrast, just before the onset of the targeting episode, B showed significantly lower rates of affinitive behaviors versus A. Reconciliation occurred in A, whereas it was absent in B. Therefore, we suggest that in B, with the decrease of baseline affinitive interactions associated with the beginning of the targeting episode, the function of postconflict reunions probably stopped working. On the whole, we found that contrasting results were probably related to different seasonal and social conditions (A: breeding/pregnancy season, characterized by higher tolerance rates; B: birth season, characterized by lower tolerance rates). Accordingly, reconciliation should be monitored throughout the different seasonal phases.     

Lactation and care for unrelated infants in forest-living ringtailed Lemurs

An adult female ringtailed lemur (Lemur catta) known not to have been pregnant showed spontaneous lactation in response to twin infants born to an unrelated female. The females had met only 7 months earlier, when they and two other unrelated adult females were released from separate locations in a forest enclosure to form a new social group. Three months after release, an adult male from an adjacent enclosure gained access to the new group for 1 day, the day of one female's estrus. No males had access to the females throughout the remainder of the breeding season. Within 2 weeks of the birth of the twins, one of the other adult females began carrying the infants frequently, typically one at a time. All three females were checked for lactation when the infants were two months old. Both their mother and the unrelated adult who had been carrying the infants were producing milk. The third adult female, who never carried either infant, had no milk. The third adult female, who never carried either infant, had no milk. This female, however, like the two maternal females, frequently attacked unfamiliar immigrating adult males when the males approached the infants. Potential implications of these observations concerning the social organization of ringtailed lemurs are discussed.