Ranging behaviour of feral cattle (Bos taurus) in Doñana National Park, S.W. Spain

The cattle of Doñana (139 individuals in four social groups in 1989) have lived under free-ranging conditions for centuries. Their ranging behaviour was analysed during a three-year period. A total of 17,603 locations corresponding to 247 different animals allowed both for the estimation of global and seasonal home ranges of individuals and social groups and for the comparison of movement patterns. Cattle ranging behaviour was not affected by human interference, and was shown to be regulated by a complex interaction of environment, individual and social factors. Habitat structure and seasonal fluctuations in abundance and distribution of resources determined general patterns of ranging behaviour: the greater the concentration of resources, the smaller the home ranges of individuals and social groups. These patterns were modified at an individual level by the sex of the animal and its reproductive status if male. Social influences on ranging behaviour were important because these implied the segregation of home ranges among dominant bulls and among social groups. As a result, there was a great variability in space use and home-range behaviour.     

Influence of the urban matrix on space use of coyotes in the Chicago metropolitan area

Expansion of the coyote’s (Canis latrans) distribution in North America has included most urban areas. Concerns for human safety have resulted in the need to understand the spatial relationship between humans and coyotes in urban landscapes. We examined the space use of coyotes with varying degrees of urban development in the Chicago metropolitan area, IL, USA, between March 2000 and December 2002. We compared home-range size, land use, and habitat use of 41 radio-collared coyotes (5 coyotes residing in developed areas, 29 in less-developed areas, and 7 in a matrix of developed and less-developed areas). The partitioning of coyotes into groups based on their level of exposure to urban development allowed us to examine if differences in use of land types by coyotes was evident in our study area. Coyotes in developed areas had home ranges twice the size of animals in less-developed areas. Nonurban habitats were used by all coyotes in the study area, while urban land was avoided. Coyotes in developed areas had large home ranges and high amounts of urban land in their range, but preferred nonurban habitat. This required the coyotes to travel through a matrix of urban land, thus encountering human activity and possibly increasing the risk of conflict with humans. However, coyotes in developed areas avoided crepuscular times when human activity was highest, suggesting that coyotes in developed areas may reduce conflicts with humans by traveling through the matrix of urban land late at night when the risk of contact with humans is lowest. Coyotes in less-developed areas were less affected by human activity at night and likely posed less risk to humans.     

Resource utilization and interspecific relations of sympatric bobcats and coyotes

We used scat analysis and radiotelemetry to characterize use of foods and habitats by sympatric bobcats and coyotes, and evaluated these in the context of spatial and temporal relationships to assess the potential for, and evidence of, interspecific competition. Bobcats and coyotes exhibited broad and overlapping diets. However, diets of the two predators differed in the relative contributions of small and large prey, with bobcats consuming relatively more rodent and lagomorph biomass and coyotes consuming relatively more ungulate biomass. Consumption among rodent prey species was highly correlated between bobcats and coyotes, indicating no evidence of prey partitioning within this group. Habitat selection by the two predators differed slightly at the landscape scale but not within home ranges. Bobcats and coyotes occupied small, overlapping home ranges, such that the likelihood of interspecific encounters (direct or indirect) was high. Bobcats displayed slight avoidance of overlapping coyote core areas during coyote reproductive seasons (winter and spring), when coyotes are typically most territorial (toward conspecifics), but displayed slight attraction during times of year when coyotes were not engaged in reproductive activities. Relative to coyotes, which were strongly nocturnal, diel activity patterns of bobcats were more diurnal and variable. However, activity patterns were not inversely correlated. Overall, these predators appeared to use resources independently and we found little evidence of negative interactions. Differences in resource use by bobcats and coyotes appeared to relate to fundamental niche differences as opposed to competition-related resource partitioning.     

Effects of hunting on the behaviour and spatial distribution of farmland birds: importance of hunting-free refuges in agricultural areas

Hunting is one of the human activities that directly affect wildlife and has received increasing attention given its socioeconomic dimensions. Most studies have been conducted on coastal and wetland areas and showed that hunting activity can greatly affect bird behaviour and distribution. Hunting-free reserves for game species are zones where birds find an area of reduced disturbance. We evaluated the effect of hunting activities on the behaviour and use of hunting-free areas of lapwings Vanellus vanellus , golden plovers Pluvialis apricaria and little bustards Tetrax tetrax in agricultural areas. We compared the habitat use and behaviour of birds on days before, during and after hunting took place. All three studied species showed strong behavioural responses to hunting activities. Hunting activity increased flight probability and time spent vigilant (higher on hunting days than just before and after a hunting day), to the detriment of resting. We also found distributional (use of hunting-free reserve) responses to hunting activities, with hunting-free reserves being used more frequently during hunting days. Thus, reserves can mitigate the disturbance caused by hunting activities, benefiting threatened species in agricultural areas. Increasing the size or number of hunting-free areas might be an important management and conservation tool to reduce the impacts of hunting activities.     

Habitat use and ranging behaviour of the red fox (Vulpes vulpes) in a Mediterranean rural area: is shelter availability a key factor?

Factors affecting activity, habitat use, and home-range size of the red fox were analysed in a highly heterogeneous rural environment. Individual differences in behaviour were used to test our hypotheses. Food habits tended to depend on food availability, which, in turn, was mainly influenced by temperature. Diet was highly heterogeneous. Insects, e.g. grasshoppers and beetles, and cultivated fruits were the staple of the diet, but no diet component stood out clearly from all others. For an opportunistic species such as the fox, habitat heterogeneity may be the main factor underlying a wide trophic niche. All foxes selected the vineyard as part of their home ranges, whereas they preferred for activity the abandoned olive-yard among the habitats of their home ranges. Human intolerance of foxes affects their pattern of activity, habitat selection, and ranging behaviour. Foxes were strongly nocturnal. Cover-rich habitats were preferred for resting and for movements in daylight. Areas under human management were mainly used at night. Selectivity was higher for resting than for activity sites. Variation in home-range size and shape can be influenced not only by the dispersion of the main food patches, but also by the location of shelters     

Spatio-temporal segregation of facultative avian scavengers at ungulate carcasses

Spatial and/or temporal segregation of resource use are mechanisms that may allow coexistence between potential competitors. Spatial and temporal patterns of carrion use were studied in the main avian scavengers of Sierra Espuña Natural Park (SE, Spain). We monitored the use of ungulate carcasses provided by hunting during winter and summer of 2005–2006. Non-breeding ravens exploited aggregated carcasses and depleted the resource in a few days while golden eagles used scattered carcasses over a longer consumption period. Moreover, non-breeding ravens exploited carcasses when golden eagles were less active. Almost all available hunter kills were exploited by golden eagles and/or ravens during winter whereas a high number of carcasses were not used during the breeding season, suggesting strong competition in a period of low food availability. Thus carrion arrangement, seasonal behaviour and competitive interactions could be affecting the foraging patterns of these two species. Interestingly, intraspecific aggregation of the smaller species and dominance of the larger one may act on the use of shared resources and trigger segregation mechanisms. Our results support that differences in the spatial and temporal patterns of resource use may allow resource partitioning between two species, thus facilitating their coexistence in sympatric areas.     

Habitat use and behaviour of European hedgehogErinaceus europaeus in a Danish rural area

HedgehogsErinaceus europaeus Linnaeus, 1758 were radio-tagged and monitored during the summer of 2001 in a Danish rural area with the objective of quantifying home ranges, nightly distances travelled, habitat use, activity patterns, day-nesting habits, and body-weight changes of the five males and five females being recorded. Males had larger home-range sizes and travelled longer nightly distances than females. The two most common habitat types within the home ranges of the hedgehogs were deciduous forest and arable land, whereas the two most frequently used habitat types were deciduous forest and grassland. No differences between the sexes were found in the proportions of different habitat types within the home ranges or in habitat use. Non-random habitat use was found; forested areas and edge habitats seemed preferred to open areas. The most frequently used day-nesting habitat was deciduous forest. Foraging was by far the most time-consuming nightly activity for both sexes. Males lost weight during the study period (May–July), whereas females gained weight. A peak in the frequency of sexual behaviour was found from late-June to mid-July. The high level of male ranging activity and the weight loss of males are interpreted as a consequence of the promiscuous mating system of hedgehogs.     

Evaluating Indirect Effects of Hunting on Mule Deer Spatial Behavior

Mule deer (Odocoileus hemionus) are widely hunted throughout western North America and are experiencing population declines across much of their range. Consequently, understanding the direct and indirect effects of hunting is important for management of mule deer populations. Managers can influence deer mortality rates through changes in hunting season length or authorized tag numbers. Little is known, however, about how hunting can affect site fidelity patterns and subsequent habitat use and movement patterns of mule deer. Understanding these patterns is especially important for adult females because changes in behavior may influence their ability to acquire resources and ultimately affect their productivity. Between 2008 and 2013, we obtained global positioning system locations for 42 adult female deer at the Starkey Experimental Forest and Range in northeast Oregon, USA, during 5-day control and treatment periods in which hunters were absent (pre-hunt), present but not actively hunting (scout and post-hunt), and actively hunting male mule deer (hunt) on the landscape. We estimated summer home ranges and 5-day use areas during pre-hunt and hunt periods and calculated overlap metrics across home ranges and use areas to assess site fidelity within and across years. We used step selection functions to evaluate whether female mule deer responded to human hunters by adjusting fine-scale habitat selection and movement patterns during the hunting season compared to the pre-hunt period. Mule deer maintained site fidelity despite disturbance by hunters with 72 ± 4% (SE) within-year overlap between summer home ranges and hunt use areas and 54 ± 7% inter-annual overlap among pre-hunt use areas and 56 ± 7% among hunt use areas. Mule deer diurnal movement rates, when hunters are active on the landscape, were higher during the hunting period versus pre-hunt or scout periods. In contrast, nocturnal movement rates, when hunters are inactive on the landscape, were similar between hunting and non-hunting periods. Additionally, during the hunt, female mule deer hourly movements increased in areas with high greenness values, indicating that mule deer spent less time in areas with more vegetative productivity. Female mule deer maintained consistent habitat selection patterns before and during hunts, selecting areas that offered more forest canopy cover and high levels of vegetative productivity. Our results indicate that deer at Starkey are adopting behavioral strategies in response to hunters by increasing their movement rates and selecting habitat in well-established ranges. Therefore, considering site fidelity behavior in management planning could provide important information about the spatial behavior of animals and potential energetic costs incurred, especially by non-target animals during hunting season. © 2020 The Wildlife Society.     

Status and diurnal activity budget of non-breeding White-headed Ducks Oxyura leucocephala in Algeria

In Algeria, the Globally Endangered White-headed Duck Oxyura leucocephala is resident throughout the year in the coastal wetlands of north-east Algeria and the Hauts Plateaux, where it occupies habitats that range from freshwater ponds and brackish marshes to hypersaline lakes. In autumn and winter, at two study sites sleeping (49% and 68%) and resting (9% and 20%) dominated, whereas feeding (7% and 10%) represented a minor proportion of, diurnal activities. There was no marked seasonal change in the activity pattern, as would be expected for a resident bird. The breeding and winter distributions of the species were similar, but seasonal and diel patterns of dispersion among habitats remain poorly known. Human encroachment on wetland habitat, habitat degradation and illegal hunting in protected areas are the major threats to the persistence of the species and probably similar threatened species in Algeria.     

Continuous remote monitoring of fish activity with restricted home ranges using radiotelemetry

Signal strength received by a fixed antenna increased as a transmitter was positioned closer to it and decreased as the transmitter was positioned further away. Depth and transmitter orientation also influenced signal strength, but these were less pronounced than the effect of distance. The variations in signal strength recorded from fixed distances, depths or orientations were low, suggesting that variation in signal strength only occurs if the transmitter is moved. The use of signal strength variation as a measure of fish activity over a 24 h period was compared with observed patterns of point habitat use of a live fish. Close correspondence of assessments of activity using signal strength variation and habitat use was observed. This study demonstrated the potential utility of radiotelemetry in association with point-in-time habitat use data to determine home-range and diel and seasonal patterns of fish activity over 24 h periods.     

Population substructure and space use of Foxe Basin polar bears

Climate change has been identified as a major driver of habitat change, particularly for sea ice-dependent species such as the polar bear (Ursus maritimus). Population structure and space use of polar bears have been challenging to quantify because of their circumpolar distribution and tendency to range over large areas. Knowledge of movement patterns, home range, and habitat is needed for conservation and management. This is the first study to examine the spatial ecology of polar bears in the Foxe Basin management unit of Nunavut, Canada. Foxe Basin is in the mid-Arctic, part of the seasonal sea ice ecoregion and it is being negatively affected by climate change. Our objectives were to examine intrapopulation spatial structure, to determine movement patterns, and to consider how polar bear movements may respond to changing sea ice habitat conditions. Hierarchical and fuzzy cluster analyses were used to assess intrapopulation spatial structure of geographic position system satellite-collared female polar bears. Seasonal and annual movement metrics (home range, movement rates, time on ice) and home-range fidelity (static and dynamic overlap) were compared to examine the influence of regional sea ice on movements. The polar bears were distributed in three spatial clusters, and there were differences in the movement metrics between clusters that may reflect sea ice habitat conditions. Within the clusters, bears moved independently of each other. Annual and seasonal home-range fidelity was observed, and the bears used two movement patterns: on-ice range residency and annual migration. We predict that home-range fidelity may decline as the spatial and temporal predictability of sea ice changes. These new findings also provide baseline information for managing and monitoring this polar bear population.     

Elephant spatial use in wet and dry savannas of southern Africa

The influence of elephants on woody vegetation cover varies from place to place. In part this may be due to the way elephants utilize space across landscapes and within their home ranges in response to the availability and distribution of food. We used location data from 18 cows at six study sites across an east to west rainfall gradient in southern Africa to test whether wet- and dry-season home-range sizes, evenness of space use within seasonal home ranges and range overlap between seasons and between years, differed between wet and dry savannas. We then tested whether the quantity, distribution and seasonal stability in vegetation productivity, a coarse measure of food for elephants, explained differences. Elephants in wet savannas had smaller wet- and dry-season home ranges and also returned to a higher proportion of previously visited grid cells between seasons and between years than elephants living in dry savannas. Wet-season home-range sizes were explained by seasonal vegetation productivity while dry-season home-range sizes were explained by heterogeneity in the distribution of vegetation productivity. The influence of the latter on dry-season home ranges differed among structural vegetation classes. Range overlap between seasons and between years was related to inter-seasonal and inter-annual stability in vegetation productivity, respectively. Evenness of elephant spatial use within home ranges did not differ between savanna types, but it was explained by seasonal vegetation productivity and heterogeneity in the distribution of vegetation productivity during the wet season. Differences in elephant spatial use patterns between wet and dry savannas according to vegetation structure and season may need to be included in the development of site-specific objectives and management approaches for African elephants.     

Behavior, Diet, and Movements of the Sulawesi Crested Black Macaque (Macaca nigra)

We present the first field study of activity budgets, diet and ranging patterns of the Sulawesi crested black macaque, Macaca nigra, one of seven macaque species endemic to Sulawesi, Indonesia. We studied three crested macaque groups, ranging in size from 50 to 97 individuals, for 18 months in the Tangkoko-DuaSudara Nature Reserve, North Sulawesi. They spent 59% of the day moving and procuring food, especially fruits, and 41% of the day resting and socializing. Their diet is composed of more than 145 species of fruit (66% of observed feeding bouts), vegetative material (2.5%), invertebrates (31.5%), and occasional vertebrate prey. Group differences were more pronounced than seasonal or diurnal differences. Specifically, the largest group moved farther during the day, moved at a faster and more uniform rate, ate less fruit, rested more, and socialized less than the smaller groups did. The largest group had the largest home range, but it included less primary forest and more disturbed habitat than the ranges of smaller groups. There are individual differences in activity budgets of adult males and females in time spent moving, resting, feeding, and socializing that may reflect differences in reproductive strategies of males versus females. The behavior of large juveniles is more similar to that of adults than to that of small juveniles. Daily movements and use of home range are correlated with diet. Macaques moved shorter distances as the proportion of time spent feeding on fruit increased, and the top four dietary items accounted for most of the variance in entry into hectare blocks of home range.     

Home-Range Use and Activity Patterns of the Red Langur (Presbytis rubicunda) in Sabangau Tropical Peat-Swamp Forest, Central Kalimantan, Indonesian Borneo

Knowledge of a species’ ranging patterns is vital for understanding its behavioral ecology and vulnerability to extinction. Given the abundance and even distribution of leaves in forested habitats, folivorous primates generally spend less time feeding; more time resting; have shorter day ranges; and require smaller home ranges than frugivorous primates. To test the influence of frugivory on ranging behavior, we established the activity budget and home-range size and use in a highly frugivorous population of the Borneo-endemic colobine, Presbytis rubicunda, within Sabangau tropical peat-swamp forest, Central Kalimantan, and examined relationships between fruit availability and ranging patterns. We collected 6848 GPS locations and 10,702 instantaneous focal behavioral scans on a single group between January and December 2011. The group had the largest home-range size recorded in genus Presbytis (kernel density estimates: mean = 108.3 ± SD 3.8 ha, N = 4 bandwidths). The annual activity budget comprised 48 ± SD 4.0% resting; 29.3 ± SD 3.9% feeding, 14.2 ± SD 2.5% traveling, and 0.4 ± SD 0.4% social behaviors. Mean monthly day-range length was the highest recorded for any folivorous primate (1645 ± SD 220.5 m/d). No significant relationships existed between ranging variables and fruit availability, and ranging behaviors did not vary significantly across seasons, potentially owing to low fluctuations in fruit availability. Our results suggest that colobine monkeys maintain larger than average ranges when high-quality food resources are available. Their extensive range requirements imply that protecting large, contiguous tracts of habitat is crucial in future conservation planning for Presbytis rubicunda.     

Effects of large-scale removal of coyotes on pronghorn and mule deer productivity and abundance

We tested the hypothesis that predation by coyotes (Canis latrans) impacts pronghorn (Antilocapra americana) and mule deer (Odocoileus hemionus) populations. We did so by examining the effects of coyote removal on pronghorn and mule deer populations within 12 large areas (>10,500 km²) located in Wyoming and Utah during 2007 and 2008. Pronghorn productivity (fawn to adult female ratio) and abundance were positively correlated with the number of coyotes removed and removal effort (hours spent hunting coyotes from aircraft) although the correlation between pronghorn productivity and removal effort was not statistically significant (P = 0.08). Mule deer productivity and abundance were not correlated with either the number of coyotes removed or removal effort. Coyote removal conducted during the winter and spring provided greater benefit than removals conducted during the prior fall or summer. Our results suggest that coyote removal conducted over large areas increases fawn survival and abundance of pronghorn but not mule deer. © 2011 The Wildlife Society.     

Landscape effects on wild boar home range size under contrasting harvest regimes in a human-dominated agro-ecosystem

Agro-ecosystems can experience elevated human-wildlife conflicts, especially crop damage. While game management often aims at reducing number to mitigate conflicts, there is on-going debate about the role of hunting disturbance in promoting game to range over wider areas, thereby potentially exacerbating conflicts. Herein, we hypothesised that landscape configuration and non-lethal disturbance modulate the response to harvest disturbance. We used an information theoretic approach to test the effects of landscape and anthropogenic variables on wild boar ranging patterns across contrasting harvest regimes. We used 164 seasonal home ranges from 95 wild boar (Sus scrofa) radio-tracked over 6 years in the Geneva Basin where two main harvest regimes coexist (day hunt and night cull). Mean seasonal 95% kernel home range size was 4.01 ± 0.20 km² (SE) and 50% core range size 0.79 ± 0.04 km², among the smallest recorded in Europe. Range sizes were larger in the day hunt area than in the night cull area, with no seasonal effect. However, when accounting for landscape variables, we demonstrate that these patterns were likely confounded by the underlying landscape configuration, and that landscape variables remain the primary drivers of wild boar ranging patterns in this human-dominated agro-ecosystem with range size best explained by a model including landscape variables only. Therefore, we recommend accounting for landscape configuration and sources of non-lethal disturbance in the design of harvest strategies when the aim is to limit wide-ranging behaviour of wild boar in order to mitigate conflicts.     

The effect of randomly altering the time and location of feeding on the behaviour of captive coyotes (Canis latrans)

Captive carnivores appear highly susceptible to the negative effects of predictable feeding routines. Current research and reviews suggest feeding unpredictably may increase species typical behaviour, but positive results have been limited to a few species. Our objectives were to determine whether randomly altering the time and location of feeding increased species typical behaviour, or affected the temporal distribution of captive coyote (Canis latrans) behaviour. We assigned 12 coyotes individually to either a predictable or an unpredictable (in time and location) feeding condition and measured behaviours in the morning and evening. Overall, mean time spent foraging, travelling, resting or standing was similar in both conditions, but coyotes in the unpredictable condition marked (118 vs. 42; P = 0.01, one-tailed) and howled (81 vs. 24; P = 0.05, one-tailed) significantly more than coyotes in the predictable condition. There were also significant temporal differences (a.m. vs. p.m.) in behaviour between conditions. Specifically, coyotes fed predictably foraged (P = 0.03) and travelled (P = 0.03) more in the morning and rested (P = 0.03) more in the evening, whereas coyotes fed unpredictably howled (62 vs. 19; P = 0.03) and stood (P = 0.05) more in the morning and foraged (P = 0.03) more in the evening. Optimum predictability for captive carnivores should be based on species-specific captive study results and relevant ecological data. Our results suggest coyote feeding regimes may be improved by including unpredictable elements nested within a predictable framework to mirror seasonal fluctuations in resources and increase species-specific behaviour in captivity.     

Intergroup relations and home range use in Verreaux's sifaka (Propithecus verreauxi)

Relationships between neighboring groups feature prominently in socioecological theory, but few empirical studies have focused on the effects of neighbors on the behavior of primates. Interactions between neighboring groups are most conspicuous during intergroup encounters, but the likelihood of encounters with neighbors can also affect ranging and activity patterns indirectly, and, as a result, behavioral patterns in areas of exclusive use may differ from those in overlap areas of adjacent home ranges. The aim of this study was to examine intergroup relations and spatial variation in home range use in Verreaux's sifaka (Propithecus verreauxi) during the annual mating season. Based on 230 hr of focal animal data collected from ten members of five adjacent groups, we found that behavioral patterns and patterns of resource utilization were not markedly different between areas of exclusive use and overlap areas of adjacent home ranges. Group cohesion tended to be higher in overlap than in core areas, but the proportion of time spent resting and foraging did not differ between these two areas. However, dominant males exhibited a higher scent-marking rate in overlap areas. Observations during intergroup encounters revealed that chases between males occurred frequently, whereas fights involving physical contact were not observed. We also found that the intergroup dominance hierarchy depended on the relative group size or the number of males in each group, with groups of lower dominance rank exhibiting a tendency to sleep proportionally more often in their core areas. The results of this study suggest that in primate species with a moderate level of intergroup aggression, such as Verreaux's sifaka, the possibility of encountering neighbors in areas of home range overlap does not exert strong influence on their behavior and resource utilization.     

Home-range size and movement patterns of Hooded Vultures Necrosyrtes monachus in southern Africa

Hooded Vulture Necrosyrtes monachus populations have declined dramatically in recent years, but we know little about their ecology. We radio-tagged four vultures in northern Botswana to gather data on animal movement and home-range patterns. Hooded Vultures were primarily sedentary at night. Hooded Vultures moved similar distances and speeds during the wet and dry season, and travelled over similar home ranges as measured using minimum convex polygons (MCP), but used much smaller core areas during the dry (breeding) season. We found significant differences in mean distances and speeds moved among different birds, and when comparing day to night, but not between the wet (non-breeding) and dry (breeding) season or by year. All of the variables we tested, including individual vulture differences, season, year and number of fixes, significantly influenced 95% MCP and kernel density estimate (KDE) home-range sizes. Hooded Vultures used significantly smaller KDE home-range sizes during the dry (breeding season) than in the wet (non-breeding) season. Hooded Vultures travelled smaller daily distances over smaller home ranges than most other vulture species for which data exist.     

Lion-tailed macaques (Macaca silenus) in a disturbed forest fragment: Activity patterns and time budget

We describe the activity patterns and time budget of a feral group of lion-tailed macaques that were confined to a disturbed forest fragment of 65 ha and compare the results with those obtained for groups in protected forests. The degraded nature of the study site was reflected in low tree densities, low specific diversity, gaps in the girth distribution of trees, and frequent disturbance by humans. The study group of 43 subjects was twice as large as lion-tailed macaque groups in protected habitats. They spent the most time ranging (34.0%), followed by foraging (23.7%), feeding (17.9%), resting (16.0%), and other activities such as social interactions (8.4%). Monthly variations are significant for all activity categories except ranging. Times spent resting and foraging are negatively correlated (r = −0.5) and show significant seasonal differences. Foraging time was highest from September to November, when key food sources such asCullenia andArtocarpus were absent or marginally available. The study group spent most time (40.4%) at canopy levels between 21 and 30 m. They spent more time each day ranging than resting or feeding and more time terrestrially compared with groups in protected forests. Large group size, poor habitat quality, and seasonal variation in food availability were the major variables affecting their time budget, and these variables accounted for differences from the time budgets of groups in protected forests.