Habitat preference in the critically endangered yellow‐tailed woolly monkey (Lagothrix flavicauda) at La Esperanza,Peru

Habitat loss is one of the main threats to wildlife. Therefore, knowledge of habitat use and preference is essential for the design of conservation strategies and identification of priority sites for the protection of endangered species. The yellow‐tailed woolly monkey (Lagothrix flavicauda Humboldt, 1812), categorized as Critically Endangered on the IUCN Red List, is endemic to montane forests in northern Peru where its habitat is greatly threatened. We assessed how habitat use and preference in L. flavicauda are linked to forest structure and composition. The study took place near La Esperanza, in the Amazonas region, Peru. Our objective was to identify characteristics of habitat most utilized by L. flavicauda to provide information that will be useful for the selection of priority sites for conservation measures. Using presence records collected from May 2013 to February 2014 for one group of L. flavicauda, we classified the study site into three different use zones: low‐use, medium‐use, and high‐use. We assessed forest structure and composition for all use zones using 0.1 ha Gentry vegetation transects. Results show high levels of variation in plant species composition across the three use zones. Plants used as food resources had considerably greater density, dominance, and ecological importance in high‐use zones. High‐use zones presented similar structure to medium‐ and low‐use zones; thus it remains difficult to assess the influence of forest structure on habitat preference. We recommend focusing conservation efforts on areas with a similar floristic composition to the high‐use zones recorded in this study and suggest utilizing key alimentation species for reforestation efforts.     

Proximal Spacing Between Individuals in a Group of Woolly Monkeys (Lagothrix lagotricha) in Tinigua National Park, Colombia

I studied proximal spacing within a group of woolly monkeys (Lagothrix lagotricha) during 7 months at Parque Nacional Tinigua, Colombia. I collected a total of 1188 instantaneous samples on focal individuals, recording the number and age/sex class of individuals that were in contact with, <2 m from, <5 m from the focal animal. The results indicate that proximate spacing reflects social affinities and is related to mother–infant relationship and social grooming. Subadult females and adult males are the sex/age classes with the lowest number of individuals in proximity. There are low proximity between adult females and between adult males and high frequencies of nearness between mother and offspring. Associations between males and females were usually low, but in some cases males showed preferences for a given female. There was a relatively gradual increase in spacing between mothers and their offspring as they became older. Old juvenile males were associated chiefly with other males—mostly subadults—whereas juvenile females maintained some proximity only to their mothers. There are also differences in spacing behavior according to different activity types.     

Long‐term monitoring of social behavior in a grouping of six female tigers (Panthera tigris)

Although literature on the reproduction of captive tigers is plentiful, there is little information on other social behaviors, particularly within large social groupings. Here we report on the long‐term monitoring of social behaviors in six female tigers (Panthera tigris). Over a period of 6 years, behavioral data were collected on spraying, vocalizations, non‐contact aggression, and contact aggression during outdoor observations, as well as nearest neighbors in the morning, as a proxy measure of affiliation. Regression analyses showed that non‐contact aggression and vocalizing were increasing, whereas spraying and social proximity were decreasing over the 6 years of the study. Paired t‐tests showed no seasonal differences in aggression, but animals had higher social proximity scores during the mornings of the colder winter months. Correlations showed that spraying was positively related to contact aggression whereas social proximity was negatively correlated with non‐contact aggression. In addition to documenting levels of contact and non‐contact aggression over time, this study showed these behaviors correlated with behaviors that were more easily scored (spraying and social proximity). Behavioral monitoring has played a key role in the management and husbandry of this large social grouping of female tigers. In addition to providing keepers with the opportunity to observe their animals in regularly scheduled sessions, behavioral monitoring has provided baseline information on social relationships in this grouping over time. If future research validates the relationship between aggression and spraying/social proximity, this can be a valuable tool for long‐term monitoring of groups of tigers. Zoo Biol 27:89–99, 2008. © 2007 Wiley‐Liss, Inc.     

Implications of Tourist–Macaque Interactions for Disease Transmission

During wildlife tourism, proximity or actual contact between people and animals may lead to a significant risk of anthropozoonotic disease transmission. In this paper, we use social network analysis, disease simulation modelling and data on animal health and behaviour to investigate such risks at a site in Morocco, where tourists come to see wild Barbary macaques (Macaca sylvanus). Measures of individual macaques’ network centrality—an index of the strength and distribution of their social relationships and thus potentially their ability to spread disease—did not show clear and consistent relationships with their time spent in close proximity to, or rate of interacting with, tourists. Disease simulation modelling indicated that while higher-ranked animals had a significantly greater ability to spread disease within the group, in absolute terms there was little difference in the size of outbreaks that different individuals were predicted to cause. We observed a high rate of physical contact and close proximity between humans and macaques, including during three periods when the macaques were coughing and sneezing heavily, highlighting the potential risk of disease transmission. We recommend that general disease prevention strategies, such as those aimed at reducing opportunities for contact between tourists and macaques, should be adopted.     

Transmission Patterns of Pinworms in Two Sympatric Congeneric Primate Species

Understanding pathogen transmission is essential to addressing the dynamics of infectious diseases in animal populations. Directly transmitted parasites spread in host populations via 1) contact with infected individuals and 2) contact with contaminated substrates. Although studies exist that support social or ranging effects on transmission, it is less clear how these factors interact. We test the hypothesis that a combination of social, ranging, diet, and intrinsic factors account for Trypanoxyuris minutus (pinworm) infections in sympatric howler species Alouatta palliata and A. pigra. We collected 211 howler fecal samples from 34 adults living in four groups, two of each species, in Tabasco (Mexico), and calculated pinworm prevalence and eggs per gram of feces (EPG). We followed each group for 80 h to determine ranging, diet, frequency of contact, and conspecific proximity. Prevalence of Trypanoxyuris minutus was high, with 82% of all individuals infected. Logistic modeling indicated that pinworm prevalence was positively associated with proximity and the proportion of group members contacted by focal individuals. Although EPG results should be interpreted cautiously owing to variable egg excretion, this index was also positively associated with proximity and the proportion of group members that were contacted, as well as with dietary diversity and use of non-tree foods. Neither intrinsic factors such as species and sex, nor group and population level variables, such as group and home range size, home range overlap, and intensity of range use, were significant predictors of pinworm infection. We conclude that both sociality and feeding behavior are key factors in infection dynamics of Trypanoxyuris minutus in sympatric Alouatta palliata and A. pigra, confirming that contact with infected conspecifics and contaminated substrates are important mechanisms for directly transmitted parasites.     

Dynamic interactions among badgers: implications for sociality and disease transmission

1. Direct interactions between individuals play an important part in the sociality of group-living animals, their mating system and disease transmission. Here, we devise a methodology to quantify relative rates of proximity interaction from radio-tracking data and highlight potential asymmetries within the contact network of a moderate-density badger population in the north-east of England. 2. We analysed radio-tracking data from four contiguous social groups, collected over a 3-year period. Dynamic interaction analysis of badger dyads was used to assess the movement of individuals in relation to the movement of others, both within and between social groups. Dyads were assessed with regard to season, sex, age and sett use pattern of the badgers involved. 3. Intragroup separation distances were significantly shorter than intergroup separation distances, and interactions between groups were rare. Within groups, individuals interacted with each other more often than expected, and interaction patterns varied significantly with season and sett use pattern. Non-mover dyads (using the main sett for day-resting on > 50% of occasions) interacted more frequently than mover dyads (using an outlier sett for day-resting on > 50% of occasions) or mover-non-mover dyads. Interactions between group members occurred most frequently in winter. 4. Of close intragroup interactions (< 50 m separation distance), 88.6% were associated with a main sett and only 4.4% with outlier setts. Non-mover dyads and non-mover-mover dyads interacted significantly more often at the main sett than mover-only dyads. These results highlight the importance of the main sett to badger sociality and support the suggestion that badger social groups are comprised of different subgroups, in our case based on differential sett use patterns. 5. Asymmetries in contact structure within a population will affect the way in which diseases are transmitted through a social network. Assessment of these networks is essential for understanding the persistence and spread of disease within populations which do not mix freely or which exhibit heterogeneities in their spatial or social behaviour.     

The time budget of a group of wild woolly monkeys (Lagothrix lagotricha)

I studied a group of woolly monkeys (Lagothrix lagotricha)for 1800 hr from early 1985 until September 1987 near the lower Apaporis river of the Vaupés Department in the eastern Colombian Amazon. The group had an activity budget of 29.9% rest, 38.8% move, 25.8% forage, and 5.5% other activities during 720 hr of observations evenly spaced throughout this year. The only significant change in these activities during the study period was social activity, which increased as a function of increased food supply. Average diurnal activities of resting and moving are negatively correlated, and each activity show significant changes during the day, while foraging show no significant change. The activity budgets of the adult males and females are not significantly different, but the activities of the juveniles and infants showed much more social activity than the adults. High moving scores may be correlated with the comparative infertility of the soils of the Caparú forest, which require increased effort to maintain the energy levels necessary for maintenance and reproduction in Lagothrix.     

Party size and diet of syntopic atelids (Ateles chamek and Lagothrix cana) in Southwestern Brazilian Amazonia.

Syntopic Alouatta seniculus, Ateles chamek and Lagothrix cana (Atelidae) were studied in southwestern Amazonia. Primate populations were first surveyed, and then the party size, diet and vertical spacing were monitored over a 5-month period. Atelids accounted for more than half the survey sightings and Lagothrix was the most abundant. Party sizes recorded for both Alouatta and Lagothrix during monitoring were significantly larger than those recorded during surveys, but no such difference was found for Ateles. Monitored parties were significantly larger in Lagothrix in comparison with either Alouatta or Ateles, as were groups of Ateles in comparison with Alouatta. Mean party size in Ateles decreased progressively during the course of the study, from 8.9 +/- 3.4 in June to 3.9 +/- 2.3 in October. Moraceae was the most important dietary resource for Ateles and Lagothrix, in terms of both feeding records and number of species exploited. There was considerable overlap in the plant taxa exploited, but some notable differences, such as the exclusive use of Hymenaea courbaril (Caesalpinaceae) by Lagothrix and of Euterpe precatoria (Arecaceae) by Ateles. As at other sites in the region, Ateles occupied significantly higher forest strata in comparison with Lagothrix. Despite the preliminary nature of the study, the results indicate a number of ecological differences between species that undoubtedly play an important role in niche separation.     

Nutritional and Health Status of Woolly Monkeys

Woolly monkeys (Lagothrix lagotricha and L. flavicauda) are threatened species in the wild and in captivity. Numerous zoological institutions have historically kept Lagothrix lagotricha spp., but only a few of them have succeeded in breeding populations. Therefore the majority of institutions that formerly kept Lagothrix lagotricha are no longer able or willing to do so. Captive populations of the species have frequent health problems, most significantly hypertension and related disorders. Researchers have conducted free-ranging dietary and behavior studies with respect to woolly monkeys, but have established no concrete link between diet or nutrients and captive health problems. The available literature we discuss indicates that researchers need to examine the link further. In addition, it is critical to the survival of the primates to be able to keep breeding populations in captivity owing to increasing natural pressures such as deforestation and hunting. Therefore, better understanding of the captive and free-ranging behavior and health parameters of the species is vital to ensure their survival and to maintain forest health and diversity. Researchers need to conduct large-scale research studies comparing the health and complete diet of individuals in the wild and captivity to resolve health problems facing the species in captivity.     

Observations on the woolly monkey (Lagothrix lagothricha) in northern Colombia

Woolly monkeys,Lagothrix lagothricha, were briefly studied in a cloud forest in Bolivar, Colombia during the dry season. Results suggest that this species is wholly arboreal, lives in multi-male heterosexual groups and does not defend territory. Woolly monkeys are folivorous and frugivorous. Social and auto-grooming are both reported. All age-sex classes except natal-coated infants and adult males indulge in play. Agonistic behaviour is very rare. There seem to be various consistent patterns of greeting behaviour and scent-marking of the environment was observed. Woolly monkeys tended to approach the observers rather than flee from them. They were observed in close proximity toCebus, Alouatta, andAteles species.     

Predicting potential distribution and identifying priority areas for conservation of the Yellow-tailed Woolly Monkey (Lagothrix flavicauda) in Peru

Species distribution models (SDMs) provide conservationist with spatial distributions estimations of priority species. Lagothrix flavicauda (Humboldt, 1812), commonly known as the Yellow-tailed Woolly Monkey, is one of the largest primates in the New World. This species is endemic to the montane forests of northern Peru, in the departments of Amazonas, San Martín, Huánuco, Junín, La Libertad, and Loreto at elevation from1,000 to 2,800 m. It is classified as “Critically Endangered” (CR) by the International Union for Conservation of Nature (IUCN) as well as by Peruvian legislation. Furthermore, it is listed in Appendix I of the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES). Research on precise estimates of its potential distribution are scare. Therefore, in this study we modeled the potential distribution area of this species in Peru, the model was generated using the MaxEnt algorithm, along with 80 georeferenced occurrence records and 28 environmental variables. The total distribution (high, moderate, and low) for L. flavicauda is 29,383.3 km², having 3,480.7 km² as high potential distribution. In effect, 22.64 % (6,648.49 km²) of the total distribution area of L. flavicauda is found within Natural Protected Areas (NPAs), with the following categories representing the largest areas of distribution: Protected Forests (1,620.41 km²), Regional Conservation Areas (1,976.79 km²), and Private Conservation Areas (1,166.55 km²). After comparing the predicted distribution with the current NPAs system, we identified new priority areas for the conservation of the species. We, therefore, believe that this study will contribute significantly to the conservation of L. flavicauda in Peru.     

Brown spider monkeys (Ateles hybridus): a model for differentiating the role of social networks and physical contact on parasite transmission dynamics

Elevated risk of disease transmission is considered a major cost of sociality, although empirical evidence supporting this idea remains scant. Variation in spatial cohesion and the occurrence of social interactions may have profound implications for patterns of interindividual parasite transmission. We used a social network approach to shed light on the importance of different aspects of group-living (i.e. within-group associations versus physical contact) on patterns of parasitism in a neotropical primate, the brown spider monkey (Ateles hybridus), which exhibits a high degree of fission–fusion subgrouping. We used daily subgroup composition records to create a ‘proximity’ network, and built a separate ‘contact’ network using social interactions involving physical contact. In the proximity network, connectivity between individuals was homogeneous, whereas the contact network highlighted high between-individual variation in the extent to which animals had physical contact with others, which correlated with an individual''s age and sex. The gastrointestinal parasite species richness of highly connected individuals was greater than that of less connected individuals in the contact network, but not in the proximity network. Our findings suggest that among brown spider monkeys, physical contact impacts the spread of several common parasites and supports the idea that pathogen transmission is one cost associated with social contact.     

Maintaining social cohesion is a more important determinant of patch residence time than maximizing food intake rate in a group-living primate,Japanese macaque (Macaca fuscata)

Animals have been assumed to employ an optimal foraging strategy (e.g., rate-maximizing strategy). In patchy food environments, intake rate within patches is positively correlated with patch quality, and declines as patches are depleted through consumption. This causes patch-leaving and determines patch residence time. In group-foraging situations, patch residence times are also affected by patch sharing. Optimal patch models for groups predict that patch residence times decrease as the number of co-feeding animals increases because of accelerated patch depletion. However, group members often depart patches without patch depletion, and their patch residence time deviates from patch models. It has been pointed out that patch residence time is also influenced by maintaining social proximity with others among group-living animals. In this study, the effects of maintaining social cohesion and that of rate-maximizing strategy on patch residence time were examined in Japanese macaques (Macaca fuscata). I hypothesized that foragers give up patches to remain in the proximity of their troop members. On the other hand, foragers may stay for a relatively long period when they do not have to abandon patches to follow the troop. In this study, intake rate and foraging effort (i.e., movement) did not change during patch residency. Macaques maintained their intake rate with only a little foraging effort. Therefore, the patches were assumed to be undepleted during patch residency. Further, patch residence time was affected by patch-leaving to maintain social proximity, but not by the intake rate. Macaques tended to stay in patches for short periods when they needed to give up patches for social proximity, and remained for long periods when they did not need to leave to keep social proximity. Patch-leaving and patch residence time that prioritize the maintenance of social cohesion may be a behavioral pattern in group-living primates.     

Risk is a Component of Social Relationships in Spider Monkeys

In group‐living animals, social relationships are the result of the interactions between two individuals over time and can provide fitness benefits to both the participants. Recently, components of social relationships were identified in Old World primates and ravens through the use of principal component analysis (PCA). We employed PCA to identify components that define the social relationships in two communities of wild spider monkeys in the Yucatán Peninsula, Mexico and investigated whether characteristics of the dyads, such as age combination, sex combination and kinship, had an effect on the components obtained. We found two components. Component 1 had high positive loadings of proximity, grooming and subgroup index, which may reflect value or compatibility. Embrace and aggression rates loaded highly on Component 2, which was labelled risk. Component 1 was affected by kinship: kin had higher scores than non‐kin. Both the components were affected by sex combination of the dyad. Male–male dyads had higher scores than female–male and female–female dyads. The results are in accordance with what is currently known about spider monkeys’ social relationships, but provide an additional perspective with the novel component of risk. In this respect, the two components are effective in capturing the ambiguous nature of male–male relationships in spider monkeys. Overall, this study offers further insight into the social patterns underlying the relationship quality in group‐living animals and the usefulness of PCA in providing an objective assessment that reflects the animals′ perspective of their social interactions.     

Performance of proximity loggers in recording intra- and inter-species interactions: a laboratory and field-based validation study

Knowledge of the way in which animals interact through social networks can help to address questions surrounding the ecological and evolutionary consequences of social organisation, and to understand and manage the spread of infectious diseases. Automated proximity loggers are increasingly being used to record interactions between animals, but the accuracy and reliability of the collected data remain largely un-assessed. Here we use laboratory and observational field data to assess the performance of these devices fitted to a herd of 32 beef cattle (Bos taurus) and nine groups of badgers (Meles meles, n?=?77) living in the surrounding woods. The distances at which loggers detected each other were found to decrease over time, potentially related to diminishing battery power that may be a function of temperature. Loggers were highly accurate in recording the identification of contacted conspecifics, but less reliable at determining contact duration. There was a tendency for extended interactions to be recorded as a series of shorter contacts. We show how data can be manipulated to correct this discrepancy and accurately reflect observed interaction patterns by combining records between any two loggers that occur within a 1 to 2 minute amalgamation window, and then removing any remaining 1 second records. We make universally applicable recommendations for the effective use of proximity loggers, to improve the validity of data arising from future studies.     

Who infects whom? Social networks and tuberculosis transmission in wild meerkats

Transmission of infectious diseases is strongly influenced by who contacts whom. Despite the global distribution of tuberculosis (TB) in free-living wild mammal populations, little is known of the mechanisms of social transmission of Mycobacterium bovis between individuals. Here, I use a network approach to examine for correlations between five distinct types of intra- and intergroup social interaction and changes in TB status of 110 wild meerkats (Suricata suricatta) in five social groups over two years. Contrary to predictions, the most socially interactive animals were not at highest risk of acquiring infection, indicating that in addition to contact frequency, the type and direction of interactions must be considered when quantifying disease risk. Within social groups, meerkats that groomed others most were more likely to become infected than individuals who received high levels of grooming. Conversely, receiving, but not initiating, aggression was associated with M. bovis infection. Incidence of intergroup roving by male meerkats was correlated with the rovers themselves subsequently testing TB-positive, suggesting a possible route for transmission of infection between social groups. Exposure time was less important than these social interactions in influencing TB risk. This study represents a novel application of social network analysis using empirical data to elucidate the role of specific interactions in the transmission of an infectious disease in a free-living wild animal population.     

Aggression and Affiliation during Social Conflict in Pigs

Social conflict is mostly studied in relation to aggression. A more integral approach, including aggressive and affiliative behaviour as well as physiology, may however give a better understanding of the animals'' experience during social conflict. The experience of social conflict may also be reflected in the spatial distribution between conspecifics. The objective was to assess the relationship between behaviour, physiology, and spatial integration in pigs (Sus scrofa) during social conflict. Hereto, 64 groups of pigs (9 wk of age) were studied in a 24 h regrouping test whereby pairs of familiar pigs were grouped with 2 unfamiliar pairs, in either barren or straw-enriched housing. Data on aggressive and affiliative behaviour, skin lesions, body weight, and haptoglobin could be summarized into three principal component analysis factors. These three factors were analysed in relation to spatial integration, i.e. inter-individual distances and lying in body contact. Pigs stayed up to 24 h after encounter in closer proximity to the familiar pig than to unfamiliar pigs. Pigs with a high factor 1 score were more inactive, gave little social nosing, had many skin lesions and a high body weight. They tended to space further away from the familiar pig (b = 1.9 cm; P = 0.08) and unfamiliar ones (b = 0.7 cm; P = 0.05). Pigs that were involved in much aggression (factor 2), and that had a strong increase in haptoglobin (factor 3), tended to be relatively most far away from unfamiliar pigs (b = 0.03 times further; P = 0.08). Results on lying in body contact were coherent with results on distances. Pigs in enriched housing spaced further apart than pigs in barren housing (P<0.001). The combined analysis of measures revealed animals that may either promote or slow down group cohesion, which may not have become clear from single parameters. This emphasizes the importance of an integral approach to social conflict.     

The time budget of a group of wild woolly monkeys (Lagothrix lagotricha)

I studied a group of woolly monkeys (Lagothrix lagotricha)for 1800 hr from early 1985 until September 1987 near the lower Apaporis river of the Vaupés Department in the eastern Colombian Amazon. The group had an activity budget of 29.9% rest, 38.8% move, 25.8% forage, and 5.5% other activities during 720 hr of observations evenly spaced throughout this year. The only significant change in these activities during the study period was social activity, which increased as a function of increased food supply. Average diurnal activities of resting and moving are negatively correlated, and each activity show significant changes during the day, while foraging show no significant change. The activity budgets of the adult males and females are not significantly different, but the activities of the juveniles and infants showed much more social activity than the adults. High moving scores may be correlated with the comparative infertility of the soils of the Caparú forest, which require increased effort to maintain the energy levels necessary for maintenance and reproduction in Lagothrix.     

Atelinae adaptations: behavioral strategies and ecological constraints.

Comparisons between the four genera that make up the Atelinae reveal two distinct behavioral patterns, one in which energy expenditure is minimized (Alouatta) and one in which energy intake is maximized (Lagothrix, Ateles, and Brachyteles). Among the atelins, Lagothrix and Ateles devote over 75% of their annual feeding time to fruit, while Brachyteles devotes between 50% and 67% of their feeding time to leaves. Pronounced seasonality in the Atlantic coastal forest inhabited by Brachyteles may be responsible for its more folivorous diet. Alouatta falls in the body size range of Lagothrix and is much smaller than Ateles and Brachyteles. Nonetheless, Alouatta is more folivorous than sympatric atelins. The atelins also share a rapid, suspensory mode of locomotion that appears to enable them to minimize travel time between widely dispersed fruit sources. Alouatta, by contrast, employs a slower, but more energetically efficient, quadrupedal locomotion. Ranging patterns among the Atelinae are consistent with both diet and locomotor abilities: Atelins travel daily distances up to 5,000 m; Alouatta ranges are much shorter. Further distinctions are evident in Atelinae grouping patterns. Alouatta remains in small cohesive groups that occupy home ranges less than 60 ha in size. Both Lagothrix and Ateles have large groups that fission to reduce the costs of intragroup feeding competition when preferred fruits occur in small patches within much larger community ranges. While greater reliance on low-energy foods such as leaves may release Brachyteles from similar competitive constraints, their tendency toward fluid grouping associations is consistent with the pursuit of a frugivorous diet.     

A Proximity-Based Social Network of a Group of Sichuan Snub-Nosed Monkeys (Rhinopithecus roxellana)

Studies of spatial proximity between individuals are important for an understanding of social structure because animals are more likely to interact with individuals in close spatial proximity. Here, we apply social network analysis to proximity data collected between 2001 and 2003 from an individually identified, provisioned, free-ranging band of Sichuan snub-nosed monkeys (Rhinopithecus roxellana) in the Qinling Mountains, central China. We aimed to quantify the social network structure and to gain insight into each individual’s position within the social network. The overall network connectivity of the study band was sparse, with a low group density of 0.17. We identified nine one-male-multifemale units (OMUs) in the study band using hierarchical cluster analysis, which confirms that this species forms a multilevel society in its natural habitat. Based on sex differences in eigenvector and betweenness centralities, adult females have more important social roles than males. Among females, lactating females scored higher betweenness and eigenvector centralities than other females. However, our results do not suggest the existence of key individual(s) in the social network of the study band. The global clustering coefficient characteristic of the band was 0.3?±?0.1, with little variation between individuals, suggesting that the removal or death of any specific individual would not significantly disrupt its general network structure. Our results also show that proximity commonly occurs among unit members, but can also occur between females of different OMUs. These observations suggest that snub-nosed monkeys have a loose-knit or fluid rather than a rigid female-bonded social system, which may be a common trend for species living in multilevel societies.