Effects of water level on sleeping-site selection and inter-group association in proboscis monkeys: why do they sleep alone inland on flooded days?

A one-male group (BE-Group) of proboscis monkeys was studied along the Menanggul River, a tributary of the Kinabatangan River, Sabah, Malaysia, from May 2005 to 2006. It has generally been assumed that proboscis monkeys only set up their sleeping sites along the riverbank; however, when more than 1 m of water covered the forest floor for more than 700 m inland from the riverbank during the seasonal flood, the BE-Group slept inside the forest. It seems that the sleeping-site selection of the BE-Group was not influenced by food availability during the flooded months because the food availability by the vegetational survey did not vary much between flooded and non-flooded months. In addition, feeding behaviors of the focal monkey in the BE-Group also did not vary much between flooded and non-flooded days. On the other hand, the water level statistically influenced the sleeping-site selection. The proboscis monkeys remained in inland forest during the flooded days because of the reduced predation threat, as terrestrial predators such as clouded leopards are prevented from foraging by deep water covering the forest floor. On non-flooded days when the BE-Group slept at the riverbank, they frequently slept close to other one-male groups on the riverside trees. Contrastingly, when the group slept inside the forest on flooded days when the water level was high, they slept away from other groups. This difference in the need for one-male groups to sleep close to each other might be attributed to the decreased predation threat during high water level in the flooded days.     

Ranging Behavior of Proboscis Monkeys in a Riverine Forest with Special Reference to Ranging in Inland Forest

We observed a unimale group (BE-Group) of proboscis monkeys (Nasalis larvatus) comprising an α-male, 6 adult females, and several immatures from May 2005 to May 2006. We followed the group for 2014 h along the Menanggul River, Sabah, Malaysia (118°30′E, 5°30′N). Observations focused mainly on ranging behavior. We determined availability and seasonal changes in plant species consumed by the members of the group by vegetation surveys in a 2.15-ha area along 200–500 m trails in the riverine forest. During the observation period, the group ranged ≤800 m from the riverbank, within a total range of 138.3 ha. The daily path length of the group ranged from 220 to 1734 m (mean, 799 m), and daily path length correlates negatively with fruit availability. The monkeys were apt to remain within a small range in fruit-abundant seasons. Because the monkeys preferred to feed on fruits of dominant plant species in the study area, their daily path length may decrease on days when they feed on fruits. The core areas of the group’s home range were along the river because the monkeys typically returned to riverside trees to sleep. The group most often used areas that were nearer the riverbank and where the availability of fruits was higher. The most frequently used grids were the ones where the group often had sleeping sites and crossed the river. Avoiding predation may be the main reason for river crossing and selecting particular sleeping sites; hence not only food availability but also the risk of predation appears to influence the ranging of the BE-Group.     

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Proboscis monkeys' (Nasalis larvatus) river crossing behavior was examined as a potential behavioral response to predation threat. N. larvatus' major predator at the Tanjung Puting National Park in Kalimantan Tengah, Indonesia, appeared to be the false gavial (Tomistoma schlegeli), a crocodilian. An adolescent female proboscis monkey was captured and drowned by a false gavial during this study. The monkeys crossed the river at its more narrow points, with several individuals or groups crossing simultaneously. Increased vigilance prior to crossing and leaping as far across the river as possible were also observed. Proboscis monkey groups often associate at their sleeping sites at the river's edge. A potential function of these associations may be to allow groups to synchronize their movement with other groups during river crossings.     

Characteristics of Night Sleeping Trees of Proboscis Monkeys (<Emphasis Type="BoldItalic">Nasalis larvatus</Emphasis>) in Sabah,Malaysia

Primates spend about half of their lives at sleeping sites, and their choice of sleeping sites may affect individual survival. We identified a total of 88 trees used by proboscis monkeys (Nasalis larvatus) as night sleeping sites on 16 nights from June to September 2008 in riverine, mangrove, and mixed mangrove–riverine forests along the Garama River, a tributary of the Klias River, in the west of Sabah, Malaysia. We recorded 11 variables for each tree, including the species, physical structure, distance from the riverbank, and connectivity with surrounding trees. We compared sleeping trees with 114 trees with ≥30 cm girth at breast height (GBH) located ≤50 m of the riverbank in 8 botanical plots (total 1 ha). Trees in the plots represented the general vegetation patterns of the study area. Choice of sleeping trees did not depend on the tree species. Although sleeping trees included trees ≤46 m from the river, those closer to riverbanks (5–35 m, n = 76) were more likely to be used as sleeping sites. Compared to the available trees, sleeping trees had larger trunks (mean±SD = 143.6 ± 56.9 cm GBH), and were taller (mean±SD = 34.3 ± 8.1 m), with greater number (median = 6; range = 12) and larger (mean±SD = 24.1 ± 15.2 cm circumference) main branches. They were also located near to other trees, with overlapping branches, creating good arboreal connectivity. Choice of sleeping trees by proboscis monkeys is likely to be related to risks of predation and injury from falling, as well as ease of social interaction and efficiency of locomotion.     

Inter-individual relationships in proboscis monkeys: a preliminary comparison with other non-human primates

This is the first report on inter-individual relationships within a one-male group of proboscis monkeys (Nasalis larvatus) based on detailed identification of individuals. From May 2005 to 2006, focal and ad libitum data of agonistic and grooming behaviour were collected in a forest along the Menanggul River, Sabah, Malaysia. During the study period, we collected over 1,968 h of focal data on the adult male and 1,539 h of focal data on the six females. Their social interactions, including agonistic and grooming behaviour, appeared to follow typical patterns reported for other colobines: the incidence of social interaction within groups is low. Of 39 agonistic events, 26 were displacement from sleeping places along the river, 6 were the α male threatening other monkeys to mediate quarrels between females and between females and juveniles, and 7 were displacement from feeding places. Although the agonistic behaviour matrix based on the 33 intra-group agonistic events (excluding events between adults and juveniles and between adults and infants) was indicative of non-significant linearity, there were some specific dominated individuals within the group of proboscis monkeys. Nonetheless, grooming behaviour among adult females within a group were not affected by the dominance hierarchy. This study also conducted initial comparisons of grooming patterns among proboscis monkeys and other primate species. On the basis of comparison of their grooming networks, similar grooming patterns among both-sex-disperse and male-philopatric/female-disperse species were detected. Because adult females in these species migrate to groups repeatedly, it may be difficult to establish the firm grooming exchange relationship for particular individuals within groups, unlike in female-philopatric/male-disperse species. However, grooming distribution patterns within groups among primate species were difficult to explain solely on the basis of their dispersal patterns. Newly immigrated females in some species including proboscis monkeys are eager to have social interactions with senior group members to improve their social position.     

Road avoidance and its energetic consequences for reptiles

Roads are one of the most widespread human‐caused habitat modifications that can increase wildlife mortality rates and alter behavior. Roads can act as barriers with variable permeability to movement and can increase distances wildlife travel to access habitats. Movement is energetically costly, and avoidance of roads could therefore impact an animal's energy budget. We tested whether reptiles avoid roads or road crossings and explored whether the energetic consequences of road avoidance decreased individual fitness. Using telemetry data from Blanding's turtles (Emydoidea blandingii; 11,658 locations of 286 turtles from 15 sites) and eastern massasaugas (Sistrurus catenatus; 1,868 locations of 49 snakes from 3 sites), we compared frequency of observed road crossings and use of road‐adjacent habitat by reptiles to expected frequencies based on simulated correlated random walks. Turtles and snakes did not avoid habitats near roads, but both species avoided road crossings. Compared with simulations, turtles made fewer crossings of paved roads with low speed limits and more crossings of paved roads with high speed limits. Snakes made fewer crossings of all road types than expected based on simulated paths. Turtles traveled longer daily distances when their home range contained roads, but the predicted energetic cost was negligible: substantially less than the cost of producing one egg. Snakes with roads in their home range did not travel further per day than snakes without roads in their home range. We found that turtles and snakes avoided crossing roads, but road avoidance is unlikely to impact fitness through energetic expenditures. Therefore, mortality from vehicle strikes remains the most significant impact of roads on reptile populations.     

Clouded leopard (<Emphasis Type="Italic">Neofelis diardi</Emphasis>) predation on proboscis monkeys (<Emphasis Type="Italic">Nasalis larvatus</Emphasis>) in Sabah,Malaysia

In this study, we have reported two direct observations of individuals from a one-male group of proboscis monkeys (Nasalis larvatus) being killed by clouded leopards (Neofelis diardi) in the riverine forest along the Menanggul river, a tributary of the Kinabatangan river in Sabah, Malaysia. One of the two individuals was an infant female and the other was a juvenile female. Based on literature reviews and the observations reported here, we suggest that clouded leopard and crocodile might be significant potential predators of proboscis monkeys of any age or sex and that predation threats elicit the monkeys' anti-predator strategies. Moreover, the observations of the monkeys' behaviour when the group is attacked by a predator suggest that the adult males in one-male groups play an important role as protectors.     

Multiple Ecological Factors Influence the Location of Proboscis Monkey (<Emphasis Type="Italic">Nasalis larvatus</Emphasis>) Sleeping Sites in West Kalimantan,Indonesia

Multiple ecological factors have been hypothesized to influence primate sleeping site selection. Testing multiple hypotheses about sleeping site selection permits examination of the relative strength of distinct ecological factors and expands our ability to understand how selection pressures influence primate sleeping behavior. Here we examine how avoidance of biting insects, thermoregulation, foraging efficiency, tree stability, and interspecific competition influence selection of sleeping sites by proboscis monkeys (Nasalis larvatus) in Indonesian Borneo. We collected data on relative insect abundance, temperature, rainfall, food availability, group size, sleeping site location, and presence of other primates for 12 mo. Using formal model comparison and information criteria, we analyzed the relative importance of these ecological factors in determining one aspect of sleeping site location: distance from the river. Our models supported the avoidance of biting insects and the foraging efficiency hypotheses. Proboscis monkeys slept further inland on nights when the abundance of sandflies was high along the river, and when less food was available along the river. Many studies suggest that primates select sleeping trees and locations to reduce predation risk; our study indicates that additional factors may also be important in determining sleeping site selection.     

Daily feeding rhythm in proboscis monkeys: a preliminary comparison with other non-human primates

In non-human primates, the daily feeding rhythm, i.e., temporal fluctuation in feeding activity across the day, has been described but has rarely received much analytical interpretation, though it may play a crucial part in understanding the adaptive significance of primate foraging strategies. This study is the first to describe the detailed daily feeding rhythm in proboscis monkeys (Nasalis larvatus) based on data collected from both riverbank and inland habitats. From May 2005 to May 2006, data on feeding behavior in a group of proboscis monkeys consisting of an alpha-male, six adult females and immatures was collected via continuous focal animal sampling technique in a forest along the Menanggul River, Sabah, Malaysia. In both the male and females, the highest peak of feeding activity was in the late afternoon at 15:00–17:00, i.e., shortly before sleeping. The differences in the feeding rhythm among the seasons appeared to reflect the time spent eating fruit and/or the availability of fruit; clearer feeding peaks were detected when the monkeys spent a relevant amount of time eating fruit, but no clear peak was detected when fruit eating was less frequent. The daily feeding rhythm was not strongly influenced by daily temperature fluctuations. When comparing the daily feeding rhythm of proboscis monkeys to that of other primates, one of the most common temporal patterns detected across primates was a feeding peak in the late afternoon, although it was impossible to demonstrate this statistically because of methodological differences among studies.     

The feeding ecology and activity budget of proboscis monkeys

A group of proboscis monkeys (Nasalis larvatus) consisting of an alpha‐male, six adult females, and several immatures was observed from May 2005–2006. We collected over 1,968 hr of focal data on the adult male and 1,539 hr of focal data on the six females in a forest along the Menanggul River, Sabah, Malaysia. Availability and seasonal changes in plant species consumed by the focal monkeys were determined by vegetation surveys carried out across an area of 2.15 ha along 200–500 m trails in riverine forest. A total of 188 plant species were consumed by the focal monkeys. The activity budget of members of our study group was 76.5% resting, 19.5% feeding, and 3.5% moving. Young leaves (65.9%) and fruits (25.9%) accounted for the majority of feeding time. Over 90% of fruit feeding involved the consumption of unripe fruits and in the majority of case both the fruit flesh and seeds were eaten. Although fruit eating was rare in some months, during other times of the year time fruit feeding exceeded the time devoted to young leaves. We found that monthly fruit availability was positively related to monthly fruit eating and feeding activity, and seasonal fluctuations in dietary diversity were significantly affected by fruit eating. These results suggest that fruit availability and fruit‐eating behaviors are key factors that influence the activity budget of proboscis monkeys. Earlier assumptions that colobine monkeys are obligate folivores do not apply well to proboscis monkeys and certain other colobines. Our findings may help contribute to a better understanding of the dietary adaptations and feeding ecology of Asian colobines. Am. J. Primatol. 71:478–492, 2009. © 2009 Wiley‐Liss, Inc.     

Sleeping sites and lodge trees of the night monkey (Aotus azarae) in Bolivia

Between 1984 and 1987, we recorded the sleeping-site and lodge tree preferences of night monkeys at the Beni Biological Station, Bolivia. We characterized the structure of sleeping-site compared lodge trees to nonlodge trees, and determined the frequency of their use. Aotus azarae used branch and liana platforms on trees of the middle strate of the forest as sleeping sites, but the lodge trees provided sparse cover. Monkeys may manipulate either natural accumulations of material or bird nests to serve as sleeping sites. The characteristics of the sleeping site and of the lodge trees may be related to protection against predators and to thermal advantages. The distribution of lodge trees appeared to be related to access to food. Activities around the sleeping site could be related to marking behavior.     

Sleeping Sites and Latrines of Spider Monkeys in Continuous and Fragmented Rainforests: Implications for Seed Dispersal and Forest Regeneration

Spider monkeys (Ateles geoffroyi) use sites composed of one or more trees for sleeping (sleeping sites and sleeping trees, respectively). Beneath these sites/trees they deposit copious amounts of dung in latrines. This behavior results in a clumped deposition pattern of seeds and nutrients that directly impacts the regeneration of tropical forests. Therefore, information on the density and spatial distribution of sleeping sites and latrines, and the characteristics (i.e., composition and structure) of sleeping trees are needed to improve our understanding of the ecological significance of spider monkeys in influencing forest composition. Moreover, since primate populations are increasingly forced to inhabit fragmented landscapes, it is important to assess if these characteristics differ between continuous and fragmented forests. We assessed this novel information from eight independent spider monkey communities in the Lacandona rainforest, Mexico: four continuous forest sites and four forest fragments. Both the density of sleeping sites and latrines did not differ between forest conditions. Latrines were uniformly distributed across sleeping sites, but the spatial distribution of sleeping sites within the areas was highly variable, being particularly clumped in forest fragments. In fact, the average inter-latrine distances were almost double in continuous forest than in fragments. Latrines were located beneath only a few tree species, and these trees were larger in diameter in continuous than fragmented forests. Because latrines may represent hotspots of seedling recruitment, our results have important ecological and conservation implications. The variation in the spatial distribution of sleeping sites across the forest indicates that spider monkeys likely create a complex seed deposition pattern in space and time. However, the use of a very few tree species for sleeping could contribute to the establishment of specific vegetation associations typical of the southeastern Mexican rainforest, such as Terminalia-Dialium, and Brosimum-Dialium.     

Sleeping site selection and presleep behavior in wild pigtailed macaques

Several factors are likely to control sleeping site selection and presleep behavior in nonhuman primates, including predation risk and location of food resources. We examined the effects of these factors on the sleeping behavior of northern pigtailed macaques (Macaca leonina). While following a troop living in the surroundings of the Visitor Center of Khao Yai National Park (Thailand), we recorded the physical characteristics and location of each sleeping site, tree, the individuals' place in the tree, posture, and behavior. We collected data for 154 nights between April 2009 and November 2010. The monkeys preferred tall sleeping trees (20.9 ± SD 4.9 m) and high sleeping places (15.8 ± SD 4.3 m), which may be an antipredator strategy. The choice of sleeping trees close to the last (146.7 ± SD 167.9 m) or to the first (150.4 ± SD 113.0 m) feeding tree of the day may save energy and decrease predation risk when monkeys are searching for food. Similarly, the choice of sleeping sites close to human settlements eases the access to human food during periods of fruit scarcity. Finally, the temporal pattern of use of sleeping sites, with a preference for four of the sleeping sites but few reuses during consecutive nights, may be a trade‐off between the need to have several sleeping sites (decreasing detection by predators and travel costs to feeding sites), and the need to sleep in well‐known sites (guaranteeing a faster escape in case of predator attack). Am. J. Primatol. 73:1222–1230, 2011. © 2011 Wiley Periodicals, Inc.     

The influence of snowfall,temperature and social relationships on sleeping clusters of Japanese monkeys during winter in Shiga Heights

We studied Japanese monkeys (Macaca fuscata) of the Shiga A₁ troop at their sleeping sites in Shiga Heights, Japan, for 41 nights during 3 winters. Monkeys chose their sleeping sites in Japanese cedars and in deciduous broad-leaved forests on non-snowing nights and in Japanese cedar forests on snowing nights. We counted 399 sleeping clusters in which 2 or more monkeys remained in physical contact through the night and 43 solitary sleeping monkeys, though monkeys did not maintain physical contact with others in the daytime. We found 397 clusters on tree branches and 2 clusters on rocks. The mean size of huddling clusters was 3.06±1.22 SD. The cluster size (3.17±1.26 SD) at lower ambient temperatures between −7 and −4°C was larger than that at higher temperatures between −2 and 4°C (cluster size 2.88±1.13 SD). Most clusters were composed of kin. Females kept close to related females in the daytime and huddled with them at night. The highest-ranking male mainly huddled with his kin and his familiar females. Other males kept farther apart from each other in the daytime, probably to avoid social conflicts. Through cold winter nights, however, such males reduced inter-individual distances and huddled with other males. Japanese monkeys appear to recognize three types of inter-individual distances: an intimate distance less than 1 m, a personal distance of 1–3 m and a social distance of 3–20 m; they change their inter-individual distances according to social and ecological circumstances.     

Multiple central place foraging by spider monkeys: travel consequences of using many sleeping sites

Summary Central place foraging models assume that animals return to a single central place such as a nest, burrow, or sleeping site. Many animals, however choose between one of a limited number of central places. Such animals can be considered Multiple Central Place Foragers (MCPF), and such a strategy could reduce overall travel costs, if the forager selected a sleeping site close to current feeding areas. We examined the selection of sleeping sites (central places) by a community of spider monkeys (Ateles geoffroyi) in Santa Rosa National Park, Costa Rica in relation to the location of their feeding areas. Spider monkeys repeatedly used 11 sleeping trees, and they tended to choose the sleeping site closest to their current feeding area. A comparison of the observed travel distances with distances predicted for a MCPF strategy, a single central place strategy, and a strategy of randomly selecting sleeping sites demonstrated (1) that the MCPF strategy entailed the lowest travel costs, and (2) that the observed travel distance was best predicted by the MCPF strategy. Deviations between the observed distance travelled and the values predicted by the MCPF model increased after a feeding site had been used for several days. This appears to result from animals sampling their home range to locate new feeding sites.     

Ranging Behavior of Proboscis Monkeys (Nasalis larvatus) in the Lower Kinabatangan, Northern Borneo

I studied the ranging behavior of proboscis monkeys (Nasalis larvatus) at two sites in the Lower Kinabatangan Region of northern Borneo. I collected data on ranging behavior via scan sampling during group follows. Groups of Nasalis larvatus had ranges overlapping those of other groups in each area. I observed no territorial behavior. Groups of Nasalis larvatus occasionally swam across the Kinabatangan River, and frequently across its tributaries. The home range size of a focal one-male group (SU1) was 220.5 ha. The group traveled farther on days when the proportion of young leaves in the diet was higher. In addition, SU1 used particular areas when they fed on flowers and fruits. Apparently, rainfall and phenology did not influence ranging patterns.     

The effect of roads on spider monkeys’ home range and mobility in a heterogeneous regenerating forest

Arboreal fauna living in tropical ecosystems may be particularly affected by roads given their dependency on forest cover and the high vulnerability of such ecosystems to changes. Over a period of 4 yr, we followed subgroups of spider monkeys living in a regenerating dry tropical forest with 8.2 km of roads within their home range. We aimed to understand whether roads shaped the home range of spider monkeys and which road features affected their movement. Only 18 percent (3 km) of the spider monkeys’ home range perimeter bordered with roads; these roads had greater habitat disparity between road sides than roads inside the home range. Although monkeys were reluctant to be close to roads, and roadside habitat contained low proportions of mature forest, spider monkeys crossed roads at 69 locations (7.5 crossings per kilometer). The main road characteristic affecting crossings was canopy opening size, with greater probability of crossing where canopy openings were smaller. Our findings support the importance of canopy opening size for road crossing of arboreal taxa, but they also indicate the relevant role roadside forest structure may have. Minimizing canopy opening size and forest disturbance along roads can facilitate the movement of arboreal fauna and preserve the important role of spider monkeys and other arboreal taxa in seed dispersal and thus the maintenance and regeneration of forest diversity.     

Effects of Habitat Degradation on Sleeping Site Choice and Use in Sahamalaza Sportive Lemurs (Lepilemur sahamalazensis)

Sleeping sites may be beneficial for animals in terms of thermoregulation, proximity to foraging sites, and protection from predators and infectious diseases. The abundance of adequate sleeping sites is thus essential for the survival of primates. We investigated microhabitats around sleeping sites, and the influence of habitat degradation on sleeping site choice and usage, in the nocturnal Sahamalaza sportive lemur, Lepilemur sahamalazensis. We used quarter point sampling (N?=?315) to describe five forest fragments and 57 sleeping sites and continuous focal animal sampling (N?=?45) to determine the diurnal activity budget, to determine whether individuals inhabiting different fragments or sleeping site types showed different levels of vigilance. Our results suggest that tall trees with large crowns, a high density of small trees, and dense canopy are particularly important for sleeping site choice. Microhabitat structure around sleeping sites did not differ between forest fragments or sleeping site types. Diameter at breast height, crown diameter, canopy cover, and bole height were similar for all sleeping trees, as were the number of lianas in trees with tree-tangle sleeping sites, and the volume of tree holes. Tree holes used as sleeping sites were most often found in dead trees of Bridelia pervilleana (50–62.5 %), whereas tree tangle sites were most often located in Sorindeia madagascariensis (20–62.5 %). Lemurs were active 5–14 % of the daytime, although they never left their sleeping sites or fed. Individuals occupying tree holes had higher levels of activity than those in tree tangles, and those in more degraded fragments were more active. Our results suggest that Sahamalaza sportive lemurs choose their sleeping sites according to specific habitat characteristics, and that factors associated with old and intact forest are likely to be crucial for their survival.     

Golden lion tamarin sleeping-site use and pre-retirement behavior during intense predation

Sleeping sites, their patterns of use, and cryptic pre-retirement behavior mitigate predation risk at sleeping sites and could influence prey fitness. We evaluated sleeping-site usage for 10 groups of golden lion tamarins (GLTs) from a population that recently suffered a substantial decline due to predation at sleeping sites. We recorded the average number of nights that groups spent at their different sleeping sites to determine whether patterns of sleeping-site use were influenced by predation risk, as measured by the rate of encounters with predators, or the availability of suitable sleeping sites, as measured by the size of a group's home range and amount of mature forest within their home range. In addition, we measured travel speed to sleeping sites and compared this speed with that recorded at other times of day. GLT groups spent more nights on average at each of their sleeping sites compared to other callitrichid species for which data are available. Predation risk and habitat characteristics were not significant predictors of how many times groups used each of their different sleeping sites. Groups significantly increased their travel speed just before entering the sleeping site. Rapid locomotion to secure tree cavities may help GLTs avoid crepuscular and nocturnal predators; however, we speculate that this strategy failed numerous GLTs in our study population during the previous decade because they used sleeping sites that were accessible to predators.     

Strategies used by bonnet macaques (Macaca radiata) to reduce predation risk while sleeping

Sleep results in a decrease in alertness, which increases an animal’s vulnerability to predation. Therefore, choice of sleeping sites would be predicted to incorporate predator-avoidance strategies. The current study, conducted in two national parks in southern India, examined the behaviors adopted by bonnet macaques (Macaca radiata) to reduce the risk of being preyed upon while sleeping. Bonnet macaques from an urban setting with a low predatory risk were included for comparison. The physical characteristics of the sleeping sites in the forest corresponded with features that were most difficult for predators to access; bonnet macaques selected emergent trees with high boles near human settlements. These trees typically overhung water. Within the canopy, individuals slept in huddled subgroups near the terminal ends of branches, preferentially selecting branches over water. Subgroups were generally composed of members of the same age and sex, which likely promoted social bonding. Adult males and females with infants selected branches higher than members of other age and sex categories. The lateral distances of individuals along branches from the main trunk were similar across demographic categories. The size of a subgroup appeared to be limited by the weight a branch could support; lateral distances were maintained by regulation of mean subgroup weight, with heavier individuals forming smaller subgroups. The urban troop slept on the top of a building. Subgroup compositions at the urban site were similar to those at the forest sites. However, subgroup size, not restricted by branch fragility, resulted in larger subgroups than those found in the forest. Our results indicate that bonnet macaques adopted a suite of behaviors that reduced their risk of being preyed upon at night by selecting sleeping sites that minimized predator encounters and by selecting the safest locations within the canopy.