Sleeping site preferences in tufted capuchin monkeys (Cebus apella nigritus)

The characteristics and availability of the sleeping sites used by a group of 27 tufted capuchin monkeys (Cebus apella nigritus) were studied during 17 months at the Iguazu National Park, Argentina. We tested different hypotheses regarding possible ultimate causes of sleeping-site selection. Most sleeping sites were located in areas of tall, mature forest. Of the 34 sleeping sites the monkeys used during 203 nights, five were more frequently used than the others (more than 20 times each, constituting 67% of the nights). Four species of tree (Peltophorum dubium, Parapiptadenia rigida, Copaifera langsdorfii and Cordia trichotoma) were the most frequently used. They constituted 82% of all the trees used, though they represent only 12% of the trees within the monkeys' home range which had a diameter at breast height (DBH) > 48.16 cm (1 SD below the mean DBH of sleeping trees). The sleeping trees share a set of characteristics not found in other trees: they are tall emergent (mean height +/- SD = 31.1+/-5.2 m) with large DBH (78.5+/-30.3 cm), they have large crown diameter (14+/-5.5 m), and they have many horizontal branches and forks. Adult females usually slept with their kin and infants, while peripheral adult males sometimes slept alone in nearby trees. We reject parasite avoidance as an adaptive explanation for the pattern of sleeping site use. Our results and those from other studies suggest that predation avoidance is a predominant factor driving sleeping site preferences. The patterns of aggregation at night and the preference for trees with low probability of shedding branches suggest that social preferences and safety from falling during windy nights may also affect sleeping tree selection. The importance of other factors, such as seeking comfort and maintaining group cohesion, was not supported by our results. Other capuchin populations show different sleeping habits which can be explained by differences in forest structure and by demographic differences.     

Links between habitat degradation,and social group size,ranging, fecundity,and parasite prevalence in the Tana River mangabey (Cercocebus galeritus)

We investigated the effects of anthropogenic habitat degradation on group size, ranging, fecundity, and parasite dynamics in four groups of the Tana River mangabey (Cercocebus galeritus). Two groups occupied a forest disturbed by human activities, while the other two occupied a forest with no human disturbance. We predicted that the groups in the disturbed forest would be smaller, travel longer distances daily, and have larger home ranges due to low food tree abundance. Consequently, these groups would have lower fecundity and higher parasite prevalence and richness (number of parasite species). We measured the abundance of food trees and anthropogenic activity in the forests, the groups' daily travel distances and home range sizes, and censused social groups over 12 months. We also analyzed fecal samples for gastrointestinal parasites from three of the groups. The disturbed forest had a lower abundance of food trees, and groups in this forest traveled longer distances, had larger home range sizes, were smaller, and had lower fecundity. The groups in the disturbed forest had higher, although not statistically significant, parasite prevalence and richness. This study contributes to a better understanding of how anthropogenic habitat change influences fecundity and parasite infections in primates. Our results also emphasize the strong influence of habitat quality in determining daily travel distance and home range size in primates. Am J Phys Anthropol, 2009. © 2009 Wiley‐Liss, Inc.     

Patterns of roost site selection and use by Southern Ground-Hornbills in north-eastern South Africa

Different habitats may be used for the needs of various aspects of an animal’s life. Southern Ground-Hornbill Bucorvus leadbeateri groups announce their presence within year-round territories by calling at dawn from their overnight roost sites. Knowledge on ground-hornbill roosting habits is limited. Groups roost in large trees, apparently close to where they end up after daily foraging. We investigated patterns of roost site selection and use for four Southern Ground-Hornbill groups in the Associated Private Nature Reserves, north-eastern South Africa, based on data from GPS-satellite transmitters. The number of roost sites used per month averaged 15.4 ± 4.7 across all groups, indicating little evidence of strong preferences for specific sites. This number was least when groups were breeding, decreasing throughout the early wet season (October–December) and was lowest during the late wet season (January–March) when actively breeding groups frequently roosted close to the nest (54–83% of roosts <1 000 m of the nest). As might be expected, the mean monthly number of nights per roost peaked during the breeding season (December–January). Riparian habitats were preferred for roosting during the breeding season, whereas disturbed areas, as well as Combretumand mopane-dominated habitats were preferred during the dry non-breeding season. Adequate large trees not only for nesting, but also for roosting, particularly in riparian habitats, may therefore be an important and potentially limiting factor for the successful reproduction of Southern Ground-Hornbills.     

Home Range, Habitat Use, and Activity of Baird's Tapir in Costa Rica1

Home range size, habitat utilization, and activity patterns of five adult Bairds tapirs (Tapirus bairdii) were studied via radiotelemetry from June 1995 to May 1996 in Corcovado National Park, Costa Rica. Estimates of 4153 animal locations were made. Home range sizes (95% minimum convex polygon) for the entire study period (wet season and dry season) averaged 125.0 ha (SD = 72.7), 94.9 ha (SD = 47.6), and 96.8 ha (SD = 51.2), respectively. Monthly home range sizes averaged 55.5 ha (SD = 32.1) and did not vary between seasons. Across the entire study, secondary forest, the most commonly used habitat type (61.3%) was utilized more than expected as predicted by availability (49.6%), while the second most used habitat, primary forest, was used (25.0%) less than expected (36.2%). The same general habitat selection pattern was observed in the dry season. Wet season habitat utilization values showed tapirs using both primary and secondary forest habitats in proportion to their availability. Diurnal and nocturnal activity levels were estimated at 20.2 and 80.4 percent, respectively. More diurnal and less nocturnal activity was observed during the wet season compared to the dry season.     

Diet and ranging behavior of the endangered Javan gibbon (Hylobates moloch) in a submontane tropical rainforest

Altitude influences forest structure and food abundance and distribution, which in turn affect primate feeding and ranging patterns. Javan gibbons (Hylobates moloch) are endemic to forests spanning a broad range of altitudes on Java, Indonesia. Most information about Javan gibbon behavior comes from studies in lowland forests, while the vast majority of wild gibbons remaining inhabit hill and lower montane forests. We studied the diets, activity patterns, and ranging behavior of three gibbon groups in hill/lower montane (950-1,100?m asl) forest in the Gunung Halimun-Salak National Park (GHSNP) from April 2008 to March 2009. The mean home range size was 37?ha and the mean daily path length was 1,180?m. The study groups spent 36% of time feeding, 41% resting, 15% traveling, 6% engaging in social behavior, and 2% in aggressive interactions. Fruit was the most important food (63% of feeding time) followed by leaves (24%), and flowers (12%). Our results suggest that Javan gibbons in higher elevation habitats have substantially larger home ranges than lowland populations, despite broad similarity in their activity budgets and diets. Conservation managers should consider the effects of altitude and habitat quality on gibbon ranging behavior when developing habitat corridors, selecting sites for translocation or reintroduction projects, and designating and managing protected areas.     

Home range structure and inter-group competition for land of Japanese macaques in evergreen and deciduous forests

The per capita home range area of Japanese macaques,Macaca fuscata, is significantly smaller in evergreen forest than in deciduous forest, though a corresponding difference in food resource utilization patterns has never been described. The present study compared the home range utilization pattern of Japanese macaques living in two habitats: the Yakushima population inhabits an evergreen forest, while the Kinkazan population inhabits a deciduous forest. We found that in the Yakushima population, (1) food density was higher; (2) inter-feeding bout sites distance was shorter; (3) daily travel distance was shorter; (4) home range size was smaller; and (5) the unit value of the main home range was higher, than in the Kinkazan population. Yakushima groups utilized a small home range area intensively, compared to Kinkazan groups. We also found that a Yakushima group shared 24% of its main home range with neighboring groups, though a Kinkazan group shared only 10% with other groups. It is supposed that food distribution affects daily ranging pattern, and ultimately the social relationships between groups in Japanese macaques.     

Home range, activity and movements of a wolf pack in central Italy

Home range, habitat use, activity and movement patterns were studied in a pack of wolves in a mountainous region of Abruzzo, central Italy from June 1986 to March 1987. The home range, estimated by the minimum convex polygon from 421 radio locations, measured 197 km² and comprised several infrastructures and areas of human presence, including four garbage dumps and two offal sites. Core areas, calculated by the harmonic mean method, were located toward the centre of the home range where human disturbance and road density were lowest but forest cover was highest. During the time-span of the study, home-range use and movement patterns suggested a marked centrality in spatial behaviour and traditionality in retreat areas year-round, both during pup-rearing season and the following months. In addition, by being essentially nocturnal, resident wolves appeared to adopt tactics of temporal segregation from people to exploit food resources safely in the proximity of human settlements. Overall activity correlated with distance travelled (r = 0.90, P << 0.001), and corresponded to cyclic nocturnal movements from retreat to feeding areas. Wolf movement rate between 20:00 and 04:00 h averaged 2.5 km/h but varied up to about 8 km/h, and daily distance travelled (= 27 km/night; range 17–38 km/night) mostly depended on the location of traditional feeding sites. Home-range configuration, habitat use, activity and movements all appeared highly integrated so as to represent the most functional compromise between avoidance of human inteference and exploitation of the available food resources.     

Home range use and habitat selection by Red deer (Cerrus elaphus) in a Sitka spruce plantation as determined by radio-tracking

The home range behaviour and habitat selection by Red deer ( Cervus elaphus L. ) in a mixed-age Sitka spruce plantation was studied from1979–82 using radio-telemetry. Females used areas with rides, young replanted and pre-thicket crops, older stands where there were checked trees, more in proportion to availability than old closed-canopy stands, open-hill ground and high-elevation newly-established forest. They used open areas more at night, dusk and dawn, and the more secluded thickets during the day. Compared to females, young males were found more in older stands, high-altitude young plantation and on open-hill ground.
Home range size (406–1008 ha for females and1062–3059 ha for males) was smaller for animals with a high proportion of favourable habitats in their range, although larger range size did not incorporate a higher total area of favourable habitat. Individual ranges overlapped.
Females used the same range from season to season and from year to year. Intensity of range usage is discussed. Males dispersed a mean distance of 15 km from their area of capture during their first or second year of age. One young male used different areas each season after dispersal. Females centred their activity in the same river catchment from season to season, but male activity centres changed from one catchment to another.
Comparison is made with the results of studies of habitat selection in the same forest using dung-accumulation techniques, and with the ranging behaviour of Red deer on open-hill ground.     

Home Range Size and Overlap of the Small Nocturnal Schlegel's Japanese Gecko (Gekko japonicus), Introduced into a City Park in Korea

Studying the home range of an organism is important in understanding its ecology. Due to being cryptic, few studies have been conducted on the home range studies of small, nocturnal geckos. We conducted radio-tracking surveys for 23 individuals to estimate the home range size and home range overlap of the Schlegel's Japanese gecko(Gekko japonicus) previously introduced into a suburban city park. Individuals were commonly found in artificial structures(buildings and accessory structures) and on nearby natural trees. Daily moved distance was positively correlated with home range size. Minimum convex polygon(MCP) home range was 97.8 m~2 for females and 99.5 m~2 for males, on average. Gekko japonicus moved farther daily distances and used wider MCP and Kernel 95 home ranges in breeding season compared to non-breeding season, while the size of Kernel 50 home range did not differ between seasons. Both daily moved distance and home range size were not significantly different between sexes. In the breeding season, MCP and Kernel 50 home ranges of each gecko overlapped with 32.4% and 13.8% of remaining geckos, respectively. Our results not only show that 1) G. japonicus uses both artificial structures and adjacent natural trees as microhabitat, but also suggest that 2) G. japonicus is non-territorial, but has a core habitat that is shared with few other individuals, and 3) the reproductive system of G. japonicus is polygamous.     

Diet, foraging, and use of space in wild golden-headed lion tamarins

Lion tamarins (Callitrichidae: Leontopithecus) are small frugi-faunivores that defend large home ranges. We describe results from the first long-term investigation of wild golden-headed lion tamarins (L. chrysomelas; GHLTs). We present data about activity budgets, daily activity cycles, diet, daily path length, home range size, home range overlap, and territorial encounters for three groups of GHLTs that were studied for 1.5-2.5 years in Una Biological Reserve, Bahia State, Brazil, an area characterized by aseasonal rainfall. We compare our results to those from other studies of lion tamarins to identify factors that may influence foraging and ranging patterns in this genus. Ripe fruit, nectar, insects, and small vertebrates were the primary components of the GHLT diet, and gums were rarely eaten. Fruit comprised the majority of plant feeding bouts, and the GHLTs ate at least 79 different species of plants from 32 families. The most common foraging sites for animal prey were epiphytic bromeliads. The GHLTs defended large home ranges averaging 123 ha, but showed strong affinities for core areas, spending 50% of their time in approximately 11% of their home range. Encounters with neighboring groups averaged two encounters every 9 days, and they were always aggressive. Data about time budgets and daily activity cycles reveal that the GHLTs spent most of their time foraging for resources or traveling between foraging sites distributed throughout their home ranges. The GHLTs spent much less time consuming exudates compared to lion tamarins in more seasonal environments. Additionally, the GHLTs had much larger home ranges than golden lion tamarins (L. rosalia), and did not engage in territorial encounters as frequently as L. rosalia. GHLT ranging patterns appear to be strongly influenced by resource acquisition and, to a lesser extent, by resource defense.     

Factors Affecting the Ranging Behavior of White-headed Langurs (<Emphasis Type="Italic">Trachypithecus leucocephalus</Emphasis>)

Ranging behavior is an important aspect of animal behavior that researchers use to investigate ecological influences on individual behavior. We studied the influence of diet, water resources, and sleeping sites on the ranging behavior of 2 groups of white-headed langurs (Trachypithecus leucocephalus) in a limestone habitat at Fusui Nature Reserve, China, between August 2007 and July 2008. During the study period, the total home range sizes for the 2 focal groups were 23.8 ha and 33.8 ha, the mean daily path lengths were 491 m and 512 m, and leaves accounted for 83.4% and 91.0% of the diet, which are well within the range of variation reported for other Trachypithecus. One focal group traveled significantly longer distances in the rainy season months than in the dry season months. This variation may be related to the seasonal difference in food availability and diet. The langurs did not use their home ranges uniformly, and 50% of their activities occurred within 11% (group 1) and 20% (group 2) of their home ranges. The most heavily used quadrats in the home ranges were located near the most frequently used sleeping sites. Moreover, the core areas (>70% of location records) of both groups’ home ranges included ≥1 permanent water pool. The langurs ventured to these pools for drinking when surface water became scarce in the dry season. These results suggest that sleeping sites and water scarcity may be significant influences on the ranging behavior of white-headed langurs in limestone habitat.     

Life on the edge: Northern bobwhite ecology at the northern periphery of their range

Over the past 40 years, Northern bobwhite (Colinus virginianus) populations have declined range-wide. The Mid-Atlantic once held the highest densities in the country and now shows some of the worst declines. Although population parameters have been quantified throughout most of the bobwhite range, Mid-Atlantic populations have been largely unstudied. To better quantify the dynamics of this declining system, we sought to not only gather annual data on home range, movement, and habitat selection, but also examine how some of these metrics might impact survival. We captured and radio-tracked 154 bobwhites between May 2006 and April 2008 on a 125 km² area of Cumberland County, New Jersey, USA. Seasonal daily movement ranged from 146 m to 158 m but several extreme movements were notable. Across seasons, grassland habitat was used in greater proportion to its availability, shrub-scrub and agriculture habitats were used equally with their availability and forests and other habitats were used less than their availability. Differences in second-order selection occurred between seasons with lower use of shrub-scrub and forest habitats and higher use of other habitat in breeding seasons. Pooled breeding season survival was 0.343, nonbreeding season survival was 0.183, and annual survival was 0.063. Although mortality was dominated by avian predators, house cat mortalities were noteworthy. Cox proportional hazard analysis revealed that risk of breeding season mortality risk was increased by longer daily movement, lower grassland use, and higher forest and other use. During the nonbreeding season, risk of mortality increased with shorter daily movement and proximity to occupied buildings and barns. This information could inform management decisions in the greater Mid-Atlantic as well as other areas of their range where they exist at very low abundances. © 2010 The Wildlife Society.     

Jaguar spacing, activity and habitat use in a seasonally flooded environment in Brazil

Between July 1980 and February 1984, six jaguars (two males, four females) were fitted with radio-collars and monitored for a cumulative total of 105 months, in the Pantanal region of southwestern Brazil. Mean home range size (minimum convex polygon) for five of them (one male, four females) was 142- 1 km². Mean home range size during the dry season was 54·3 km², whereas in the wet season it was significantly smaller, 12·8 km². Mean home range overlap of the four females was 42%. Use of gallery forest and forest patches exceeded the availability of these habitat types in the animals' home ranges, whereas open forest and grassland were used less than expected on the basis of their availability. Mean distance moved between locations on consecutive days was 2·4 km (0.2-10·4 km). The mean one-day movement of the male jaguar was significantly (P<0·001) larger than that of the females. Mean distance travelled by all animals during one-day intervals in the dry season was significantly greater (P<0·001) than that travelled in other months. Jaguars were more active during daytime than night-time (P<0·001). Overall level of activity for the wet season did not differ from that of the dry season. With their larger body size, jaguars in the Pantanal may require more food than jaguars in other areas of the species' range. This, in addition to differences in prey availability and the annual availability of dry land, appear to influence the difference in home range size between jaguar populations.     

Spatial and temporal patterns in the activity of European moles

 Using ratio-tracking data obtained at three sites, we assessed the effects of season and of neighbour avoidance on the activity and patterns of home range use by European moles (Talpa europaea). The home ranges of non-breeding male and female moles did not differ significantly in size, and averaged 2324 m² (minimum convex polygon). Although overlap between ranges was small (an average of 12.8% of each range being shared with neighbours and an average of only 3.3% of 2×2 m grid cells were shared with an individual neighbour, ranges were not oriented to avoid neighbours. Non-breeding male/female neighbours tended to share more of their grid cells (3.9±5.7%) mean ±SD than did neighbours of the same sex (male:male 1.2±0.95%; female:female 1.1±1.3%), but there was no significant difference in overlap between any combination of sex pairings. On average, each mole spent only 0.9% of its time within 6 m of another mole, and only 3 out of 46 dyads showed evidence of being attracted to each other; there was no evidence from the simultaneous movement patterns of neighbouring moles that they avoided each other. Although moles tended to return to the same part of their range at the same time on successive days, there was also some indication of gradual changes in the spatial pattern of daily home range use. Moles had a triphasic pattern of activity, but this became tetraphasic under drought conditions. There were significant differences between sites, but not between sexes, in sleeping behaviour and activity patterns. These differences could be related to seasonal differences in soil moisture and thus probably to prey renewal rates. We conclude that in our sites, the activity patterns and movements of moles depend on the temporal and spatial dispersion of food, rather than on short-term interactions between the movements of neighbours. Received: 13 January 1996 / Accepted: 26 June 1996     

Home-range use and intergroup encounters in western gorillas (Gorilla g. gorilla) at Lossi forest, North Congo

I present data on home-range use and types of intergroup encounters for one group (Apollo) of western gorillas (Gorilla gorilla gorilla) from a new study site in the Republic of Congo. The total home-range size of the focal group, which I calculated by superimposing a 100 m x 100 m grid over the mapped daily path traveled, was 11 km2. The majority (73%) of the group's home range was used exclusively, although at the periphery it overlapped with the ranges of three other groups. Most encounters (86%) with other groups (n = 14) took place in the periphery of the home range, and appeared to involve access to fruit trees. The focal group silverback's encounters with solitary silverbacks occurred throughout the focal group's home range, did not involve access to fruit, and typically resulted in aggressive or avoidance behavior. The focal group silverback's response to other group males was more varied: it included tolerance (64%), avoidance (14%), and aggression (21%), and was dependent upon the identity of the extragroup male. The focal group exhibited an unusual form of tolerant behavior toward some other groups by occasionally forming "nesting supergroups" (two groups nested together overnight at distances of 30-50 m). The western gorillas at Lossi were somewhat fluid in their grouping. Subgrouping and supergrouping occurred, although more infrequently than reported previously, and with a new twist: subgrouping did not necessarily require a silverback's presence. I stress the need for intraspecific comparisons and more complete data sets on western gorilla social organization.     

Effects of Seasonal Water Scarcity on the Ranging Behavior of Eulemur fulvus rufus

The climate of western Madagascar is characterized by a long (8–9 mo) dry season during which small rivers run dry, so that most animals are dependent on access to a few permanent water holes. We studied the effects of water scarcity at the end of the dry season on the ranging behavior of 4 groups of redfronted lemurs (Eulemur fulvus rufus) living at different distances from the Kirindy Riverbed in the Kirindy/CFPF forest northeast of Morondava. Using radio-collars, we located 2 resident groups with permanent home ranges near the river, and 2 non-resident groups and followed them for 60 days. We obtained a GPS reading every 30 min and later analyzed them with GIS ArcView to determine the size and location of each group's (core) home range. One group resided in direct proximity to the riverbed with several water holes and made regular short trips (mean 330 m) to drink. A second resident group had no direct access to water and made regular trips to the same water hole at a mean distance of 590 m from their home range. One non-resident group had a center of activity about 1300 m from the nearest water hole, to which they traveled on a non-daily basis. The other non-resident group established a temporary binuclear home range with one center of activity near the riverbed, about 2900 m away from the other center of activity. Thus, redfronted lemurs use different drinking and ranging tactics as a function of the distance of their core areas from the nearest water source.     

Patterns of range use,range defense,and intergroup spacing in moustached tamarin monkeys (Saguinus mystax)

In this paper we examine patterns of group spacing and habitat utilization in neighboring groups of marked free-ranging moustached tamarin monkeys (Saguinus mystax) inhabiting Padre Isla, a small island in the Amazon Basin of northeastern Peru, and describe the set of behavioral mechanisms used by tamarins to maintain the spatial isolation and social integrity of individual groups. Specifically, we address a series of questions regarding the importance of resource defense, mate defense, and territorial defense in intergroup interactions. From June through November 1990, we recorded 67 intergroup interactions involving members of our two main study groups. These interactions occurred at a rate of .14/observation hour and were of two general types. Vocal battles averaged 18 min in duration and were characterized by a series of high frequency, short syllable, long calls that were exchanged between groups separated by distances of greater than 25 m. Aggressive encounters averaged 26 min in duration and involved visual contact, alarm calls, scent marking, and sequences of chases and retreats. Intergroup confrontations did not cluster around the perimeter of a group's home range, and there was no evidence that moustached tamarins patrolled range borders. Our data indicate that 35% of aggressive encounters occurred in the vicinity of major feeding trees. Priority access to these sites is likely to have an important influence on tamarin foraging success. Mate defense and the exploration of new breeding opportunities also appear to be important functions of intergroup conflicts. Not only did the frequency of aggressive encounters increase during breeding periods, but three-fourths of all observed copulations occurred during or within 30 min of an encounter. Given the high degree of reproductive competition reported among tamarin females and the time and energy group members devote to intergroup aggression, maintaining access to a stable home range and the resources contained within that range appear to be critical functions of moustached tamarin social interactions.     

Ranging behavior of the François’ langur (<Emphasis Type="Italic">Trachypithecus francoisi</Emphasis>) in the Fusui Nature Reserve,China

Data on activity budgets and ranging patterns were collected from March to December 2001 for one group of François’ langurs (Trachypithecus francoisi) inhabiting a forested part of the Fusui Nature Reserve, Guangxi province, China. Our results indicate that the total size of the home range of the study group during the study period was 19 ha. The majority of their activities (52%) occurred within a small area, 22%, of their home range, and was concentrated in or near quadrats containing their sleeping sites, which may reduce the time and energetic cost of travel. The extent of the ranging behavior varied between months, with the smallest, 7 ha, recorded in July and the largest, 13.5 ha, in November. There was no significant difference between seasons. The monthly mean daily path lengths varied from 341 to 577 m. The daily path lengths showed significant seasonal changes: the path lengths were longer during the dry season than in the rainy season, which may be related to the scarcity of preferred food resources during the dry season.     

Demography, range use, and behavior in black lemurs (Eulemur macaco macaco) at Ampasikely, northwest Madagascar

We studied a black lemur population over a 2-year period (1992-1993) and 8 years later (2000) in a 50-ha secondary forest in northwest Madagascar. All of the animals were marked to investigate population dynamics and seasonal variation in ranging and behavior, and new data on black lemurs were obtained. Our data on demographic characteristics were expanded to include other forest sites and contrasted with those collected in other Eulemur macaco macaco field studies, in relation to human activity and the presence of introduced and cultivated plant species. Density is affected by deforestation and hunting. Group size and home range depend on the composition of the forest and probably food patches. Sex ratio at birth varies according to the number of females per group, a result that fits the local resource competition model. Groups are multimale-multifemale, and adult females form the core of the groups. Reproductive parameters indicate sharply defined seasonal breeding, a high female reproductive rate, and birth synchrony. Changes in group composition reveal male and female juvenile dispersal, male transfer between groups at the time of mating, and adult female transfer and group fission when groups exceed a critical size. At mating and birth, intergroup agonistic encounters occurred at home-range boundaries, and larger groups were dominant over smaller groups. Patterns of intragroup interactions suggest that males compete for access to groups of females during the mating season, and that females may compete for food resources during the birth season. Our study also reports female social dominance and lack of sexual weight dimorphism in this species.