Spontaneous tool use in captive, free-ranging golden lion tamarins (Leontopithecus rosalia rosalia)

Two types of tool use were observed in eight captive, free-ranging golden lion tamarins (Leontopithecus rosalia rosalia). All eight individuals used twigs and/or radio collar antennae to pry bark from trees and probe crevices, presumably for invertebrates. Three individuals used tools for grooming. In two animals, antennae were used as grooming tools while the third individual used a stick while grooming. The complexity of the free-ranging environment may have played a role in the expression of tool use behavior in these animals, as tool use has never been observed in captive tamarins living in traditional enclosures or wild tamarins. Social transmission may be one of the mechanisms responsible for the acquisition of tool use — six of the eight tool users resided in two social groups, and the only two individuals that used antennae as grooming tools were a bonded pair. These are the first published observations of tool use by golden lion tamarins or any callitrichid in a non-experimental setting and provide further data supporting the theory of a link between extractive foraging and tool use.     

Habitat Exploitation by Free-ranging Saguinus niger in Eastern Amazonia

We monitored the foraging behavior of the members of a group of black-handed tamarins (Saguinus niger) at a site in eastern Amazonia. Their diet was frugivorous-insectivorous, but also included pod exudate of Parkia pendula. The focal group used all 3 types of forest—primary, logged, and secondary— in all months, but differentially between seasons. In the dry season, tamarins spent more than half of activity time in primary forest and less than a third in secondary forest whereas during the wet season, the proportions were reversed. Data on resource abundance indicated that the shift in habitat preference is related to a seasonal change in the distribution of fruit sources, with a larger number of species and individuals fruiting during the wet season. We recorded no such variation in the abundance of arthropods. While using a larger area, including more secondary forest, during the wet season, the group traveled significantly shorter distances each day, reflecting the availability of a larger number of fruit sources. Overall, the results reemphasize the ecological and behavioral flexibility of Saguinus niger and their ability to cope with habitat disturbance.     

Wild geladas (Theropithecus gelada) in crops—more than in pasture areas—reduce aggression and affiliation

Human–primate interfaces are expanding and, despite recent studies on primates from peri-urban environments, little research exists on the impact of agriculture and/or pasture areas on primate social behavior and health. We assessed how crop/pasture areas potentially alter social behavior and health of wild geladas (Theropithecus gelada) frequenting the unprotected area of Kundi (Ethiopia). We predicted that compared to pasture areas, crop areas (i) would be more challenging for geladas (prediction 1) and (ii) would have a greater impact on both aggressive and affiliative behavior, by reducing grooming time and enhancing competition (prediction 2). During January–May 2019 and December 2019–February 2020, we collected data (via scan, focal animal sampling, and video analyses) on direct human disturbance, external signs of pathologies and social behavior of 140 individuals from 14 one-male units and two all-male units. Animals experienced the highest level of human disturbance in crop areas (in line with prediction 1). Individuals from the groups preferentially frequenting crop areas showed the highest prevalence of external signs of pathologies consistent with chemical and biological contamination (alopecia/abnormally swollen parts). We collected 48 fecal samples. Samples from frequent crop users contained the highest rates of parasitic elements/gram (egg/larva/oocyst/cyst) from Entamoeba histolytica/dispar, a parasite common in human settlements of the Amhara region. In crop areas, subjects spent less time grooming but engaged in lower rates of intense aggression (in partial agreement with prediction 2). We speculate that the reduction in social behavior may be a tactic adopted by geladas to minimize the likelihood of detection and maximize food intake while foraging in crops.

     

Personality predicts ectoparasite abundance in an asocial sciurid

Parasitism is a consequence of complex interactions between host, parasite, and their shared environment, and host behavior can influence parasite risk. Animal personality (i.e., consistent behavioral differences that are repeatable across time and context) can influence parasitism with more explorative individuals typically hosting greater parasite loads. Host “sociality” is known to impact parasite risk with more social individuals typically at higher risk of acquiring or transmitting parasites, but other behaviors could also be important. We quantified personality in least chipmunks (Tamias minimus), including repeatability of behavioral traits, and determined whether these personality traits affected ectoparasite prevalence and abundance. We measured personality using standardized hole‐board tests and quantified ectoparasitism of 39 least chipmunks over 2 years at a site in southeastern Manitoba, Canada. We found that activity and exploration were repeatable within the context of the hole‐board test for least chipmunks, which suggests that these traits reflect personality. More exploratory individuals hosted a greater abundance of ectoparasites compared to less exploratory individuals. Our results are consistent with past studies implicating personality as a factor in host–parasite dynamics and suggest that exploration may be an important behavioral correlate of parasite acquisition.     

Vocal Repertoire of Golden-backed Uakaris (<Emphasis Type="Italic">Cacajao melanocephalus</Emphasis>): Call Structure and Context

The study of vocal behavior can reveal important aspects of how and why a species communicates in relation to ecological and social challenges. We here focus on vocal communication in golden-backed uakaris (Cacajao melanocephalus), diurnal, pitheciine monkeys that exhibit fission-fusion social organization and typically inhabit dense forests that limit the potential for visual communication. Moreover, the species spends little time engaged in tactile or olfactory communication, e.g., social grooming and scent marking, respectively. Hence, vocalizations may be very important for the coordination of social organization in these monkeys. We 1) categorized golden-backed uakari vocalizations, 2) ascertained their behavioral context, and 3) investigated whether golden-backed uakari calls can encode information about the signaler. We observed the monkeys during 2 wet seasons in the flooded igapó forest of Jaú National Park, Brazil. We showed that golden-backed uakaris have 9 call types in their vocal repertoire, all distinguishable by ear and from analysis of spectrograms. Some calls, e.g., play-specific calls, were used only in particular behavioral contexts, and by individuals of specific age, whereas others were emitted under a range of situations. The structure of the loud tchó call varied among individuals, and according to behavioral context, i.e., whether individuals were foraging/feeding, traveling, or performing agonistic interactions. This knowledge of the species’ vocal repertoire is valuable for surveying the monkeys acoustically in habitats where visual surveys are difficult.     

A multiyear survey of helminths from wild saddleback (Leontocebus weddelli) and emperor (Saguinus imperator) tamarins

The establishment of baseline data on parasites from wild primates is essential to understand how changes in habitat or climatic disturbances will impact parasite–host relationships. In nature, multiparasitic infections of primates usually fluctuate temporally and seasonally, implying that the acquisition of reliable data must occur over time. Individual parasite infection data from two wild populations of New World primates, the saddleback (Leontocebus weddelli) and emperor (Saguinus imperator) tamarin, were collected over 3 years to establish baseline levels of helminth prevalence and parasite species richness (PSR). Secondarily, we explored variation in parasite prevalence across age and sex classes, test nonrandom associations of parasite co‐occurrence, and assess the relationship between group size and PSR. From 288 fecal samples across 105 individuals (71 saddleback and 34 emperor tamarins), 10 parasite taxa were identified by light microscopy following centrifugation and ethyl‐acetate sedimentation. Of these taxa, none were host‐specific, Dicrocoeliidae and Cestoda prevalences differed between host species, Prosthenorchis and Strongylida were the most prevalent. Host age was positively associated with Prosthenorchis ova and filariform larva, but negatively with cestode and the Rhabditoidea ova. We detected no differences between expected and observed levels of co‐infection, nor between group size and parasite species richness over 30 group‐years. Logistic models of individual infection status did not identify a sex bias; however, age and species predicted the presence of four and three parasite taxa, respectively, with saddleback tamarins exhibiting higher PSR. Now that we have reliable baseline data for future monitoring of these populations, next steps involve the molecular characterization of these parasites, and exploration of linkages with health parameters.     

Grooming relationships between breeding females and adult group members in cooperatively breeding moustached tamarins (Saguinus mystax)

Grooming is the most common form of affiliative behavior in primates that apart from hygienic and hedonistic benefits offers important social benefits for the performing individuals. This study examined grooming behavior in a cooperatively breeding primate species, characterized by single female breeding per group, polyandrous matings, dizygotic twinning, delayed offspring dispersal, and intensive helping behavior. In this system, breeding females profit from the presence of helpers but also helpers profit from staying in a group and assisting in infant care due to the accumulation of direct and indirect fitness benefits. We examined grooming relationships of breeding females with three classes of partners (breeding males, potentially breeding males, (sub)adult non-breeding offspring) during three reproductive phases (post-partum ovarian inactivity, ovarian activity, pregnancy) in two groups of wild moustached tamarins (Saguinus mystax). We investigated whether grooming can be used to regulate group size by either "pay-for-help" or "pay-to-stay" mechanisms. Grooming of breeding females with breeding males and non-breeding offspring was more intense and more balanced than with potentially breeding males, and most grooming occurred during the breeding females' pregnancies. Grooming was skewed toward more investment by the breeding females with breeding males during the phases of ovarian activity, and with potentially breeding males during pregnancies. Our results suggest that grooming might be a mechanism used by female moustached tamarins to induce mate association with the breeding male, and to induce certain individuals to stay in the group and help with infant care.     

Unraveling the dispersal patterns and the social drivers of natal emigration of a cooperative breeding mammal,the golden lion tamarin

The study of the social drivers of animal dispersal is key to understanding the evolution of social systems. Among the social drivers of natal emigration, the conspecific attraction, aggressive eviction, and reduced social integration hypotheses predict that sexually mature individuals who receive more aggressive behavior and are engaged in less affiliative interactions are more likely to disperse. Few reports have explored these proximate factors affecting emigration in cooperatively breeding species, particularly of Neotropical primates. In this study, we investigated the dispersal patterns and tested the social drivers of natal emigration in the golden lion tamarin (Leontopithecus rosalia) — an endangered species inhabiting Atlantic rainforests fragments in Brazil. We used behavioral and demographic data collected during 7 years from 68 groups of tamarins inhabiting 20 forest fragments. Our analyses from the 160 dispersing individuals showed that dispersal success is higher for males and for those engaged in parallel dispersal, but that males and females use different strategies to enhance their dispersal success, males immigrate into established groups while females form new groups. We did not find high levels of agonistic behavior among group members before natal emigration. Instead we found that conspecific attraction drives natal emigration in both sexes, while additionally the low level of affiliative interactions within the natal group triggers male emigration. We discuss natal emigration in the broader perspective of the cooperative breeding system and the implications of these findings for the conservation of the species.     

Variable infant care contributions in cooperatively breeding groups of wild saddleback tamarins

Among non-human primates, alloparental infant care is most extensive in callitrichines, and is thought to be particularly costly for tamarins whose helpers may suffer increased energy expenditure, weight loss, and reduced feeding time and mobility. The costs and benefits of infant care likely vary among group members yet very few wild studies have investigated variable infant care contributions. We studied infant care over an 8-month period in four wild groups of saddleback tamarins in Bolivia to evaluate: (a) what forms of infant care are provided, by whom, and when, (b) how individuals adjust their behavior (activity, vigilance, height) while caring for infants, and (c) whether individuals differ in their infant care contributions. We found that infant carrying, food sharing, and grooming varied among groups, and immigrant males—those who joined the group after infants were conceived—participated less in infant care compared to resident males. Adult tamarins fed less, rested more, and increased vigilance while carrying infants. Although we did not detect changes in overall activity budgets between prepartum and postpartum periods, tamarins spent more time scanning their environments postpartum, potentially reflecting increased predation risk to both carriers and infants during this period. Our study provides the first quantitative data on the timing and amount of infant carrying, grooming, and food transfer contributed by all individuals within and among multiple wild groups, filling a critical knowledge gap about the factors affecting infant care, and highlighting evolutionary hypotheses for cooperative breeding in tamarins.     

A field study of the ecology and behavior of the black-mantle tamarin (Saguinus nigricollis)

The author made an ecological study of the black-mantle tamarin (Saguinus nigricollis) in the primary forest in the basin of the River Caquetá in the upper Amazon in Colombia for 88 days during a study period from August 1975 to February 1976. The tamarin group repeats three activity patterns — feeding, resting, and moving — regularly in a day. In their feeding activity, the tamarins preferred to eat small insect and resin. They showed a special liking for large grasshoppers, 6–8 cm long, and spent much of the day to capture them. Moving in search of the grasshoppers, which are hard to capture in the forest, is closely related to the tamarin group's daily nomadism. From this evidence it can be concluded that the behavior which forms the basis of the tamarins' daily activity is the hunting of grasshoppers. In the transition time from the feeding activity, principally the hunting of grasshoppers, to the resting activity, a fruit eating behavior was always observed. At this time the tamarins ate well-ripened sweet fruit intensively for a short time. This behavior is thought to have the effect, in one respect, of easing strained relations arising among group members during the grasshopper hunting activity. It was also observed that mother tamarins shared their hard-to-capture grasshoppers with their offspring although the latter had already become independent of their mothers. In general, the tamarins lived in a small and compact group consisting of an adult male, an adult female, and their offspring. Some of these small groups had amicable interrelationships and often moved together, whereas other groups kept aloof from each other. These facts indicate that, besides the groups as the social units, large-groups comprising some of these unit groups moving together, exist in the social structure of the tamarins, and form the upper structure of the social unit. See the note added to the end of this article.     

Social behavior of wild moustached tamarins,Saguinus mystax,at the Estación Biológica Quebrada Blanco,Peruvian Amazonia

The social behavior in two small, polyandrous groups of moustached tamarins, Saguinus mystax, was studied at the Estación Biológica Quebrada Blanco in northeastern Peru. Allogrooming was the most obvious type of social interaction. Grooming relations were characterized by a strong asymmetry. In both groups, one adult male contributed about 70% of all allogrooming given (in terms of time spent grooming) and allocated his allogrooming evenly to all other group members. Grooming relations between the second adult male and other group members were less intensive. Intragroup agonistic interactions were infrequent, and most of them occurred in the context of feeding on small food resources (fruit, exudate) and during intergroup encounters. Sexual behavior was rare, but at least in one group a tendency existed for the principal groomer to be somewhat more active than the other male. Results are compared with data from other field studies on callitrichid social behavior. Although the label polyandrous correctly describes the demographic structure of many groups, it is not clear whether it is sufficient for understanding the patterning of social relations and thus the social organization of tamarins. © 1996 Wiley-Liss, Inc.     

Locomotor behavior and feeding ecology of the panamanian tamarin (Saguinus oedipus geoffroyi,callitrichidae, primates)

The Callitrichidae are a family of New World primates that exhibit a complex of behavioral and morphological characters reputedly similar to those of tree squirrels of the genus Sciurus.In particular, the locomotor behavior of tamarins and marmosets has been described as “squirrel-like. ” In this paper I describe a field investigation of the locomotor behavior and ecology of the Panamanian tamarin (Saguinus oedipus geoffroyi)and the redtailed squirrel (Sciurus granatensis).From January through August 1978, a total of 1200 hr was spent observing freeranging populations of tamarins and tree squirrels in an area of secondary forest on the Pacific Coast of Panama. Observations were made using an instantaneous time sampling technique. Individual activity records were collected on focal animals and recorded at 2.5-min intervals throughout the day. The following information was collected: (a) nature and structure of the substrate exploited, (b) activities on these supports and/or types of resources procured, and (c) body posture involved in the use of these supports. Data presented indicate major differences in substrate preference and positional behavior in tamarins and tree squirrels. Unlike squirrels, tamarins avoid vertical and sharply inclined supports during travel. Movements through the canopy is accomplished by a series of long leaps which begin and end on thin terminal supports. However, the Panamanian tamarin spent numerous hours clinging to large vertical trunks while feeding on plant exudate. Gums comprise 23.O% of the noninsect portion of the tamarin diet. The relationships between small body size, claw-like nails, substrate preference, and positional behavior are discussed. Claw-like nails enable this primate to exploit a food resource that would otherwise be inaccessible. The interrelationship between environment, behaviour, and morphology provides a frameworks from which to understand callitrichid adaptations. These adaptations are not convergent with those of the sciurid rodents. An earlier draft of this paper was presented at the 48th Annual Meeting of the American Association of Physical Anthropologists, San Francisco, April 1979.     

Use of Social and Ecological Information in Tamarin Foraging Decisions

A major consequence of group living is that foragers may rely on social information in addition to ecological information to locate feeding sites. Although conspecifics can provide cues as to the spatial location of food patches, individual foraging decisions also must include some assessment of the likelihood of obtaining access to a resource other group members seek. This likelihood differs in the 2 models generally proposed to explain intragroup social foraging: the information-sharing model and the producer-scrounger model. We conducted an experimental field study on wild groups of emperor (Saguinus imperator) and saddleback (S. fuscicollis) tamarins to determine the foraging strategies adopted by individual group members and their relationship to social rank, food intake, and the ability to use ecological and social information in making intra-patch foraging decisions. Individual tamarins applied different behavioral strategies compatible with a finder-joiner paradigm to solve foraging problems. About half of the individuals in each study group initiated 74%–90% of all food searches and acted as finders. Most alpha individuals adopted a joiner strategy by monitoring the activities of others' to obtain a reward. The individual arriving first at a reward platform enjoyed a finder's advantage. Despite differences in search effort, both finders and joiners presented similar abilities in learning to associate ecological cues with the presence of food rewards at our experimental feeding stations. We conclude that within a group foraging context, tamarins integrate social and ecological information in decision-making.     

Effects of forest fragmentation on the lekking behavior of White-throated Manakins in Central Amazonia

Forest fragmentation can affect various aspects of population dynamics, but few investigators have assessed possible effects on the behavior of a species. Loss of habitat may limit population recruitment and abundance, which may alter breeding dynamics in forest remnants. We examined the lekking behavior of White-throated Manakins (Corapipo gutturalis) in a fragmented landscape to determine if forest fragmentation affected the spatial distribution of display courts and male behavior at courts. We captured and observed males at 19 courts located in 11 primary forests of different sizes in forest habitats of the Biological Dynamics of Forest Fragments Project area, an experimentally fragmented landscape located in the central Brazilian Amazon, and estimated their spatial distribution as the distance to the nearest court in the landscape. We quantified habitat loss using the proportion of forest cover surrounding courts and their distances to forest edges. No courts were detected in 1-ha forest fragments, suggesting direct effects from habitat loss following fragmentation that affected connectivity and thus recruitment and persistence of courts in the smallest fragments. The spatial distribution of display courts in forests larger than 10 ha remained unaltered, compared to display courts in continuous forests, but adult males were less numerous on courts with a higher percentage of forest cover and they displayed less on courts closer to forest edges. The spatial distribution of courts also contributed to variation in male social behavior, with more juvenile males present and adult males displaying at lower rates at more isolated courts. Although White-throated Manakins are locally common, the observed behavioral changes in response to habitat loss may affect their population dynamics. Our results show the importance of assessing behavioral changes in conservation programs and, in particular, of including biologically relevant measures of habitat loss in addressing its possible effects on species persistence in fragmented landscapes.     

Whom to Groom and for What? Patterns of Grooming in Female Barbary Macaques (Macaca sylvanus)

Grooming is one of the most conspicuous social interactions among nonhuman primates. The selection of grooming partners can provide important clues about factors relevant for the distribution of grooming within a social group. We analyzed grooming behavior among 17 semi-free ranging female Barbary macaques (Macaca sylvanus). We tested whether grooming is related to kinship, rank and friendship. Furthermore, we tested whether grooming is reciprocated or exchanged for rank related benefits (i.e. lower aggression and increased tolerance whilst feeding). We found that in general grooming was reciprocally exchanged, directed up the hierarchy and at the same time affected by friendship and kinship. Grooming was more frequent among individuals with higher friendship values as well as amongst related individuals. We also divided our data set on the basis of rank difference and tested if different power asymmetries between individuals affected the tendency to exchange grooming for rank related benefits and grooming reciprocation. In support of our initial hypothesis our results show that the reciprocation of grooming was a significant predictor of grooming interactions between individuals of similar rank, but not between those individuals more distantly separated in the social hierarchy. However, we did not find any evidence for grooming being exchanged for rank related benefits in either data set. Our results, together with previously published studies, illustrate the behavioral flexibility of macaques. It is clear that multiple studies of the same species are necessary to gather the data required for the solid comparative studies needed to shed light on patterns of grooming behavior in primates.     

The Effect of Crowding on the Social Behavior of the Cooperatively Breeding Cotton-Top Tamarins (<Emphasis Type="Italic">Saguinus oedipus</Emphasis>)

Several studies show that primates exhibit coping strategies to buffer the effect of increased social tension under crowded conditions. This is the first study to assess the effect of crowding on the social behavior of a cooperative breeding primate. We studied the effect of repeated short-term crowding on the social behavior of 20 cotton-top tamarins (10 breeders, 10 adult–subadult offspring) belonging to 2 large and 3 small groups. We compared scratching and rates of social behavior between 2 conditions: large-outdoor (42.0 m² × 3.3 m) and small-indoor (3.5 m² × 2.2 m) enclosures. Our results indicate that scratching increases in breeders and offspring during crowding. However, whereas the social behavior of breeders in small groups was not affected by crowding, breeders in large groups (n = 4) showed increased rates of severe aggression with their offspring during crowding. Offspring of large groups showed a nonsignificant trend to increase mild aggression, and an increased percentage of time in contact and grooming during crowding. This augmentation of grooming occurred in offspring–breeder dyads and also between offspring. These results suggest that helpers from large groups use coping strategies when social tension increases.     

Grooming behavior of spider monkeys (Ateles geoffroyi) on barro colorado Island,Panama

Quantitative grooming data are presented for free-ranging black-handed spider monkeys (Ateles geoffroyi) on Barro Colorado Island, Republic of Panama. A total of 126 grooming sessions was recorded, with an average session length of 2.0 min (range, 0.1 to 10.0 min). Grooming was an infrequent behavior; on average, individuals allocated only 2.5% of their daily activity to grooming. Two daily peaks of grooming activity were observed, one near midday and another in the late aftermoon between 1600 and 1700. Adult females groomed most frequently, followed by males and then juveniles. Juveniles were the most frequent recipients of grooming, followed by females and then males. Individual preferences were observed primarily between mother-offspring, male-male, and juvenile-male grooming partners in this male-bonded fission-fusion, species. Grooming interactions reflect many of the social characteristics of spider monkey societies: intraclass grouping preferences, long period of juvenile dependence, male philopatry, and female dispersal.     

Monkeys in the Middle: Parasite Transmission through the Social Network of a Wild Primate

In wildlife populations, group-living is thought to increase the probability of parasite transmission because contact rates increase at high host densities. Physical contact, such as social grooming, is an important component of group structure, but it can also increase the risk of exposure to infection for individuals because it provides a mechanism for transmission of potentially pathogenic organisms. Living in groups can also create variation in susceptibility to infection among individuals because circulating levels of immunosuppressive hormones like glucocorticoids often depend on an individual’s position within the group’s social structure. Yet, little is known about the relative roles of socially mediated exposure versus susceptibility in parasite transmission among free-living animal groups. To address this issue, we investigate the relationship between host dominance hierarchy and nematode parasite transmission among females in a wild group of Japanese macaques (Macaca fuscata yakui). We use social network analysis to describe each individual female’s position within the grooming network in relation to dominance rank and relative levels of infection. Our results suggest that the number of directly-transmitted parasite species infecting each female, and the relative amount of transmission stages that one of these species sheds in faeces, both increase with dominance rank. Female centrality within the network, which shows positive associations with dominance hierarchy, is also positively associated with infection by certain parasite species, suggesting that the measured rank-bias in transmission may reflect variation in exposure rather than susceptibility. This is supported by the lack of a clear relationship between rank and faecal cortisol, as an indicator of stress, in a subset of these females. Thus, socially mediated exposure appears to be important for direct transmission of nematode parasites, lending support to the idea that a classical fitness trade-off inherent to living in groups can exist.     

Anthropogenic impacts on Costa Rican bat parasitism are sex specific

While anthropogenic impacts on parasitism of wildlife are receiving growing attention, whether these impacts vary in a sex‐specific manner remains little explored. Differences between the sexes in the effect of parasites, linked to anthropogenic activity, could lead to uneven sex ratios and higher population endangerment. We sampled 1108 individual bats in 18 different sites across an agricultural mosaic landscape in southern Costa Rica to investigate the relationships between anthropogenic impacts (deforestation and reductions in host species richness) and bat fly ectoparasitism of 35 species of Neotropical bats. Although female and male bat assemblages were similar across the deforestation gradient, bat fly assemblages tracked their hosts closely only on female bats. We found that in female hosts, parasite abundance per bat decreased with increasing bat species richness, while in male hosts, parasite abundance increased. We hypothesize the differences in the parasite–disturbance relationship are due to differences in roosting behavior between the sexes. We report a sex‐specific parasite–disturbance relationship and argue that sex differences in anthropogenic impacts on wildlife parasitism could impact long‐term population health and survival.     

An ethogram forLemur fulvus

As a result of over 2000 hours of observation of two captiveLemur fulvus groups, their social behavior can now be provisionally described and categorized. Characteristic postures forL. fulvus include the normal standing or walking posture, often accompanied by low-pitched contact grunting, and the tightly curled resting posture. Disturbances in the vicinity of the group may lead to gradational changes in individual postures, culminating in the mob display. Greeting behaviors include passing without interaction, the sniff greeting, tactile greetings, and anogenital marking of one lemur by another. Allogrooming, mutual grooming, and reciprocal grooming may follow initial contact.L. fulvus individuals scent-mark spaces and objects, doing so at an increased rate during the mating season. Throughout most of the year little behavior occurs which can accurately be labelled ‘aggressive’. True agonistic behavior has only been observed during certain phases of the annual cycle: birth season, mating season, and the time of sexual maturation of juvenile animals. Sexual behavior is also rarely observed. Maternal behavior is interesting not only in terms of mother-infant interactions but also with regard to changes in mother-other interactions over time. Finally, play behavior can be distinguished from other kinds of behavior with similar behavioral components by looking at itsGestalt. Most play can be classified as either primarily locomotor/manipulative or primarily social.