Locomotion and feeding postures of spider and howling monkeys: field study and evolutionary interpretation

Field observations demonstrate clear differences in locomotion and feeding postures between spider monkeys (Ateles) and howling monkeys (Alouatta). When feeding, Ateles employs sitting postures approximately half the time, and a variety of suspensory postures using the tail the other half. Ateles moves quadrupedally during 52% of locomotion, by tail-arm suspension 25%, and various mixed support-suspensory modes the remainder. Tail-arm suspension is practiced more rapidly on thinner supports, and on more negatively inclined supports than is quadrupedal movement. Howlers do not locomote by tail-arm suspension: movement is almost entirely quadrupedal and is slower than that of spider monkeys. The positional behavior of spider monkeys fits closely recent views of major adaptive changes in hominoid evolution emphasizing brachiation and speed during travel. Howler locomotion and also tissue composition appear related to diet and digestive mechanisms.     

Locomotor behavior of wild orangutans (Pongo pygmaeus wurmbii) in disturbed peat swamp forest, Sabangau, Central Kalimantan, Indonesia

This study examined the locomotor behavior of wild Bornean orangutans (P. p. wurmbii) in an area of disturbed peat swamp forest (Sabangau Catchment, Indonesia) in relation to the height in the canopy, age-sex class, behavior (feeding or traveling), and the number of supports used to bear body mass. Backward elimination log-linear modeling was employed to expose the main influences on orangutan locomotion. Our results showed that the most important distinctions with regard to locomotion were between suspensory and compressive, or, orthograde (vertical trunk) and pronograde (horizontal trunk) behavior. Whether orangutans were traveling or feeding had the most important influence on locomotion whereby compressive locomotion had a strong association with feeding, suspensory locomotion had a strong association with travel in the peripheral strata using multiple supports, whereas vertical climb/descent and oscillation showed a strong association with travel on single supports in the core stratum. In contrast to theoretical predictions on positional behavior and body size, age-sex category had a limited influence on locomotion. The study revealed that torso orthograde suspension dominates orangutan locomotion, concurring with previous studies in dipterocarp forest. But, orangutans in the Sabangau exhibited substantially higher frequencies of oscillatory locomotion than observed at other sites, suggesting this behavior confers particular benefits for traversing the highly compliant arboreal environment typical of disturbed peat swamp forest. In addition, torso pronograde suspensory locomotion was observed at much lower levels than in the Sumatran species. Together these results highlight the necessity for further examination of differences between species, which control for habitat.     

The ecological role of the prehensile tail in white-faced capuchins (Cebus capucinus).

Prehensile tails appear to have evolved at least twice in platyrrhine evolution. In the atelines, the tail is relatively long and possesses a bare area on the distal part of its ventral surface that is covered with der-matoglyphs and richly innervated with Meissner's corpuscles. In contrast, the prehensile tail of Cebus is relatively short, fully haired, and lacks specialized tactile receptors. Little is currently known regarding tail function in capuchins, and whether their prehensile tail serves a greater role in feeding or traveling. In this paper we examine patterns of positional behavior, substrate preference, and tail use in wild white-faced capuchins (Cebus capucinus) inhabiting a wet tropical forest in northeastern Costa Rica. Observational data were collected over the course of 3 months on adult capuchins using an instantaneous focal animal time sampling technique. Differences in the frequency and context of tail use, and the estimated amount of weight support provided by the tail relative to other appendages during feeding/foraging and traveling were used as measures of the ecological role of this specialized organ in capuchin positional behavior. During travel, quadrupedal walking, leaping, and climbing dominated the capuchin positional repertoire. The capuchin tail provided support in only 13.3% of travel and was principally employed during below branch locomotor activities. In contrast, tail-assisted postures accounted for 40.6% of all feeding and foraging records and occurred primarily in two contexts. The tail was used to suspend the individual below a branch while feeding, as well as to provide leverage and weight support in above-branch postures associated with the extraction of prey from difficult to search substrates. A comparison of tail use in Cebus, with published data on the atelines indicates that both taxa possess a grasping tail that is capable of supporting the animal's full body weight. In capuchins and howling monkeys, the tail appears to be used more frequently and serves a greater weight-bearing role during feeding than during traveling. In Ateles, and possibly Brachyteles, and Lagothrix, however, the prehensile tail serves a dual role in both feeding and forelimb suspensory locomotion. Additional relationships between white-faced capuchin feeding, positional behavior, extractive foraging techniques, and prehensile tail use are discussed.     

Seasonal variation in the positional behavior of red howling monkeys (Alouatta seniculus)

Recent studies on the positional behavior of primates reveal that significant seasonal variation occurs in both locomotion and postures that is related to changes in diet and foraging techniques. Howling monkeys (genusAlouatta), which also have a seasonally varied diet, are predicted to have correspondingly varied positional behaviors. Two groups of red howling monkeys were studied in a primary rain forest in French Guinana during the dry and wet seasons. During the dry season, when howler diet is based mainly on leaves, howlers traveled more frequently by quadrupedal walking on large supports, a mode of progression that is probably inexpensive energetically and relatively stable. During feeding, quadrupedal and tripedal stand contributed considerably, a posture probably associated with the equal distribution of leaves within a tree crown. In contrast, during the wet season, when fruit was abundant, howlers fed very frequently by sitting on large supports, probably because fruit consumption required more time for special manipulation. However, most seasonal changes in feeding postures, and in travel and feeding locomotion, were difficult to associate directly with dietary shifts. These behavioral changes may be more highly correlated with slight modifications in microhabitat use (horizontal and vertical daily ranges, similar and alternative arboreal pathways) that are not considered in this paper.     

Positional behavior of the white uakari (Cacajao calvus calvus)

The positional behavior and habitat use of a group of white uakaries (Cacajao calvus calvus) was observed for 6 weeks in the dry season at Lake Teiú, Brazil. Data are presented for feeding, traveling, and resting activities. The most common feeding posture is sit, followed by stand. Cacajao frequently exhibits locomotor behaviors while in feeding trees, using pronograde clamber and quadrupedal walk. The most frequently used locomotor behaviors in travel are quadrupedal walk, leap, and pronograde clamber. Quadrupedal run and drop also figure importantly in the behavioral repertoire. The most frequent resting posture was sit, followed by ventral lie. Compared to representative members of the other pitheciin genera, Pithecia and Chiropotes, Cacajao engages in more locomotion while feeding, and uses more pedal suspension. While traveling, pronograde clamber and drop are more frequently used by Cacajao. Multiple, deformable supports are used more by Cacajao than by the other pitheciins throughout all activities. Overall, the positional behavior of Cacajao is more similar to that of Chiropotes than of Pithecia. Cacajao's behavioral solutions to the problems of balance imposed by its greatly reduced tail are discussed. © 1996 Wiley-Liss, Inc.     

Positional behavior of female bornean orangutans (Pongo pygmaeus)

Observational data were collected on the positional behavior of habituated adult female orangutans in the rain forest of the Kutai National Park, East Kalimantan, Indonesia. Results revealed the following about locomotion during travel: movement was concentrated in the understory and lower main canopy; and brachiation (without grasping by the feet) accounted for 11% of travel distance, quadrupedalism for 12%, vertical climbing for 18%, tree-swaying for 7%, and clambering for 51%. In climbing and clambering, the animal was orthograde and employed forelimb suspension with a mixture of hindlimb suspension and support. Thus suspension by the forelimbs occurred in at least 80% of travel. Locomotion in feeding trees resembled that during travel but with more climbing and less brachiation. Feeding was distributed more evenly among canopy levels than was travel, and use of postures (by time) included sitting 50%, suspension with the body vertical 11%, and suspension by hand and foot with the body horizontal 36%. The traditional explanation of the evolution of the distinctive hominoid postcranium stresses brachiation. More recently it has been proposed that climbing, broadly defined and partly equivalent to clambering in this study, is the most significant behavior selecting for morphology. The biomechanical similarity of brachiation and the orthograde clambering of orangutans precludes the present results from resolving the issue for the evolution of Pongo. The orangutan is by far the largest mammal that travels in forest canopy, and a consideration of the ways that its positional behavior solves problems posed by habitat structure, particularly the tapering of branches and gaps between trees, indicates that suspensory capacities have been essential in permitting the evolution and maintenance of its great body size.     

Positional behavior,selective use of habitat substrate and associated non-positional behavior in free-rangingColobus guereza (Rüppel, 1835)

Positional behavior (posture and locomotion), associated social/maintenance behaviors and selective utilization of available habitat structure were recorded for free-rangingColobus guereza occupying a remant montane forest near Limuru, Kenya. This research design focuses on relationships between anatomy, behavior, and the environment and provides a framework for interpreting adaptations inColobus. Quantitative analyses show daily activity patterns. These monkeys spend most of their time in above branch postural activities, primarily in sitting postures. Locomotion accounts for less than one-fifth of positional behavior. In both postural and locomotor sequences, the forelimbs may be placed above the shoulder complex, often in abducted positions. Suspensory positional behavior is rare. Most activity occurs within Zone II, flexible branches of medium diameter and less than 45° in grade. Selective use of the substrate and preferred positional modes are related to variables such as social/maintenance activities, weight distribution of the link system and substrate configuration. These data suggest that, (a) black and white colobus do not exhibit behavior associated with so-called “semibrachiation” and (b) bio-behavioral adaptations involving foraging and feeding strategies may account, in part, for observed morphological differences among Old World monkeys.     

Diets of some French guianan primates: Food composition and food choices

We studied food choices of black spider monkeys (Ateles paniscus) and red howlers (Alouatta seniculus) in an undisturbed tropical forest of French Guiana for 6 months in the rainy season. We made additional observations on tufted capuchins (Cebus apella) and examined the differences and similarities in feeding behavior with respect to the plant specific composition of the habitat and the biochemical characteristics of their food. Capuchins and spider monkeys mainly fed on ripe fruit pulp, to which they added invertebrates (capuchins) or young leaves (spider monkeys); their plant diet was more varied than that of howlers, which contained approximately equal proportions of ripe fruits and young leaves of many species, with large monthly variations in these food categories. BothAteles andAlouatta tended to feed preferentially on abundant plant species, with a large overlap in their fruit choices. The former species included a high proportion of soluble sugars and a low proportion of protein in its diet compared, to howlers. Leaves of several species selected byAlouatta reacted positively when screened for alkaloids and phenolic compounds. Besides specific metabolic requirements and adaptations to deriving nutrients from distinct food types, we hypothesize that the ability to taste sugars, which varies among primates, affects the range of foods appearing to be palatable and, consequently, contributes to the differentiation of feeding niches.     

Positional behavior ofCebuella pygmaea in Yasuni National Park, Ecuador

In terms of positional behavior, the small-bodied callitrichids are distinguished from other anthropoids by their ability to leap between and to cling to large vertical supports, feeding occasionally or frequently on sap and bark insects. In this paper, I studied the positional behavior of a group of pygmy marmosets in a hilly wet tropical forest in Yasuni National Park, Ecuador. During traveling, the animals used quadrupedal walk/bound and leaping mostly on small horizontal supports. During foraging, quadrupedal walk/bound and clambering dominated. The main foraging postures were stand and cantilever, occurring mainly on small horizontal lianas. During feeding, scansorial locomotion was used very frequently. Claw clinging was the dominant feeding posture. Furthermore, large vertical lianas and tree boles were the most frequent feeding supports. These observations would suggest that scansorial locomotion, vertical clinging, and vertical leaping are most likelynot part of the same form function complex. Vertical leaps appear to be associated with moving in the lower parts of the forest. On the other hand, scansorial locomotion and vertical clinging appear to be related to feeding on the sap of tree boles and large lianas in the lower parts of the forest.     

Comparisons of Suspensory Behaviors Among <Emphasis Type="Italic">Pygathrix cinerea</Emphasis>, <Emphasis Type="Italic">P. nemaeus</Emphasis>, and <Emphasis Type="Italic">Nomascus leucogenys</Emphasis> in Cuc Phuong National Park,Vietnam

In our study at the Endangered Primate Rescue Center of Cuc Phuong National Park, Vietnam, we aimed first to assemble a positional behavioral profile of captive gray-shanked (Pygathrix cinerea) and red-shanked (P. nemaeus) doucs that relates to the use of forelimb suspensory postures and arm-swinging locomotion. The profile is of interest because researchers have documented that red-shanked doucs more frequently use suspensory postures and locomotions than other colobines do. We confirmed that red-shanked doucs commonly use suspensory positional behaviors and also that gray-shanked doucs use suspensory behaviors at similar or even higher frequencies than those of red-shanked doucs. Our second goal was to assemble a preliminary kinematic profile of suspensory locomotion in Pygathrix within the context of the arm-swinging locomotion exhibited by northern white-cheeked gibbons, Nomascus leucogenys. Mean forelimb angles at initial contact and release of arm-swinging behaviors were remarkably consistent among gibbons and doucs despite the fact that gibbons typically used more continuous brachiation. Doucs also exhibit a greater range of forelimb angles than gibbons do. In addition, trunk orientation tends to be less vertical at initial contact for doucs than for gibbons, perhaps owing to the frequent use of quadrupedal sequences directly before or after forelimb suspension. Our behavioral and kinematic analyses add to the emerging realization that Pygathrix is capable of, and frequently expresses, a range of suspensory positional behaviors, including brachiation.     

Feeding and Resting Postures of Wild Northern Muriquis (Brachyteles hypoxanthus)

Increased body size in Brachyteles has been regarded as an important evolutionary adaptation that allowed a greater reliance on leaves compared to other more frugivorous Atelidae, but its association with muriqui positional behavior and substrate use is still unknown. Here, we present original data on the feeding and resting postures of the northern muriqui (Brachyteles hypoxanthus) and evaluate predictions about the relationships between body size, postural behavior, and substrate use derived from previously published data for other atelids (e.g. Alouatta, Ateles, and Lagothrix). The study was undertaken from August 2002 to July 2003 on a large group of well‐habituated muriquis inhabiting the Reserva Particular do Patrimônio Natural – Felíciano Miguel Abdala in Minas Gerais, Brazil. Consistent with our predictions, we found that B. hypoxanthus was highly suspensory during postural feeding (60.9%) and commonly used tail–hind limb suspension/horizontal tripod (38.0%) or tail–forelimb/hind limb suspension (21.4%). However, although tail‐suspensory postures permitted the muriquis to use the terminal canopy and small‐sized substrates, these areas were also accessed via tail‐assisted above‐branch postural behaviors involving multiple substrates. Unexpectedly, tail‐suspensory postures were found to be frequently associated with large substrates, tree trunks, and the understory. We suggest that Brachyteles’ ability to access food resources from all areas of a feeding tree and from tree crowns at different canopy levels may account for their ability to efficiently exploit food resources in seasonal disturbed forest fragments of the Brazilian Atlantic Forest today. Am. J. Primatol. 75:74‐87, 2013. © 2012 Wiley Periodicals, Inc.     

Suspensory locomotion of Lagothrix lagothricha and Ateles belzebuth in Yasuní National Park,Ecuador

A comparative field study of the locomotion of woolly monkeys (Lagothrix lagothricha) and spider monkeys (Ateles belzebuth) in undisturbed rainforest of northeastern Ecuador reveals substantial differences in their use of suspensory modes. Ateles performed both more brachiation (by forelimbs and tail, with trunk rotation), and forelimb swing (similar to brachiation, but without trunk rotation) than Lagothrix. In contrast, in Lagothrix 20% of suspensory movement was by pronograde forelimb swing, which resembles forelimb swing except that the body is held in a pronograde orientation due to the tail and/or feet intermittently grasping behind the trailing forelimb. Ateles never exhibited this mode. Both brachiation and forelimb swing by Ateles were more dynamic than in Lagothrix, consisting of higher proportions of full-stride bouts (versus single-step). Both species used smaller supports for suspensory than for quadrupedal locomotion, and Ateles used both smaller and larger supports for suspension than did Lagothrix. Analysis of support inclination shows that both species tended to perform more above-support movement on horizontal supports and more below-support (suspensory) movement from oblique supports. Our attempt to elucidate the aspects of canopy structure that favor suspension suggests the need for additional kinds of observational data, focusing on the "immediate structural context" of positional events.     

Arboreal locomotor and postural behaviour of European red squirrels (<Emphasis Type="Italic">Sciurus vulgaris</Emphasis> L.) in northern Greece

The study of the locomotion and postures of arboreal squirrels may provide important contextual information on the evolution of the morphology and ecology of sciurids. In this context, we studied the positional behaviour and habitat use of four adult European red squirrels (Sciurus vulgaris L.) in a mixed coniferous forest in northern Greece. Our results show that, during the study period, S. vulgaris extensively used the forest canopy and the terminal branch zone. The use of small and medium supports of all orientations was also particularly frequent. The positional profile of the species was characterized by the dominance of quadrupedal, clawed and airborne locomotion along with seated and standing postures. Quadrupedalism and sitting appeared to promote terminal branch use for food access and manipulation, while claw climbing favored vertical ranging and retreat to trees after terrestrial foraging. Finally, leaping reduced energetic costs during travelling between food sites within the relatively dispersed forest. These results and those of previous research on the positional behaviour of other squirrels reveal several trends related to body size, arboreal or gliding habits and tropical or temperate forest distribution and contribute to the understanding of evolutionary novelty in multiple levels within the sciurid radiation.     

Tool use in wild spider monkeys (<Emphasis Type="Italic">Ateles geoffroyi</Emphasis>)

Tool use has been observed in a variety of primate species, including both New and Old World monkeys. However, such reports mainly address the most prodigious tool users and frequently limit discussions of tool-using behavior to a foraging framework. Here, we present observations of novel and spontaneous tool use in wild black-handed spider monkeys (Ateles geoffroyi), where female spider monkeys used detached sticks in a self-directed manner. We introduce factors to explain Ateles tool-using abilities and limitations, and encourage the synthesis of relevant research in order to gain insight into the cognitive abilities of spider monkeys and the evolution of tool-using behaviors in primates.     

Foraging,Food Choice,and Food Processing by Sympatric Ripe-Fruit Specialists: <Emphasis Type="Italic">Lagothrix lagotricha poeppigii</Emphasis> and <Emphasis Type="Italic">Ateles belzebuth belzebuth</Emphasis>

Studies of interspecific competition and niche separation have formed some of the seminal works of ecology. I conducted an 18-mo study comparing the feeding ecologies of 2 sympatric, closely-related ripe-fruit specialists, Humboldt's woolly monkeys (Lagothrix lagotricha poeppigii), and the white-bellied spider monkeys (Ateles belzebuth belzebuth) in Amazonian Ecuador. Woolly monkeys in the terra firme forest live at roughly triple the density of spider monkeys (31 versus 11.5 animals/km²). Woolly monkeys spend 17% of their time foraging, while spider monkeys spend only 1% of their time foraging. Spider monkeys alone fed on soil and termitaria, which are rich in phosphorus. Woolly monkeys are not hard-fruit specialists. Their fruit diet is significantly more diverse than that of spider monkeys. Dietary overlap between the 2 species is high, yet each specializes to some degree on a different set of fruit resources. Woolly monkeys visit more food sources per unit of time, feed lower in the canopy, visit more small food patches, and prey on more seeds. Spider monkeys feed on fewer, richer food sources and are more than twice as likely to return to a particular fruit source than woolly monkeys are. Spider monkeys maximize fruit pulp intake, carrying more intact seeds in their guts, while woolly monkeys minimize seed bulk swallowed through more careful food processing. Surprisingly, several preferred spider monkey foods with high fat content and large seeds are avoided by woolly monkeys. I outline the different ecological dimensions involved in niche separation between the 2 species and discuss the possible impetus for their evolutionary divergence.     

Program of the fifty-sixth annual meeting of the American Association of Physical Anthropologists to be held in New York City Vista International Hotel April 2–5, 1987

Male orangutans (Pongo pygmaeus) weigh about twice as much as females. Observations in northern Sumatra of adults of both sexes (one male, three females) feeding in the same trees reveal the effects of body size dimorphism on feeding behavior: The male tended to use larger branches than the females, and to employ above-branch postures (sitting and standing) with greater frequency. The females employed suspensory under-branch postures more often. When feeding techniques were variable, the male tended to pull in branches to detach food with the mouth, whereas the female plucked more fruit by hand. By controlling for postcranial morphology and habitat structure, these results provide the first rigorous quantitative test of predictions about the effects of body size on primate positional behavior, and raise further questions about sexual diethism in feeding postural behavior of primates of varying absolute size.     

Positional behavior and substrate use in wild adult bearded capuchin monkeys (Sapajus libidinosus)

Natural selection for positional behavior (posture and locomotion) has at least partially driven the evolution of anatomical form and function in the order Primates. Examination of bipedal behaviors associated with daily activity patterns, foraging, and terrestrial habitat use in nonhuman primates, particularly those that adopt bipedal postures and use bipedal locomotion, allows us to refine hypotheses concerning the evolution of bipedalism in humans. This study describes the positional behavior of wild bearded capuchins (Sapajus libidinosus), a species that is known for its use of terrestrial substrates and its habitual use of stones as tools. Here, we test the association of terrestrial substrate use with bipedal posture and locomotion, and the influence of sex (which co‐varies with body mass in adults of this species) on positional behavior and substrate use. Behavior and location of 16 wild adult bearded capuchins from two groups were sampled systematically at 15 s intervals for 2 min periods for 1 year (10,244 samples). Despite their different body masses, adult males (average 3.5 kg) and females (average 2.1 kg) in this study did not differ substantially in their positional behaviors, postures, or use of substrates for particular activities. The monkeys used terrestrial substrates in 27% of samples. Bipedal postures and behaviors, while not a prominent feature of their behavior, occurred in different forms on the two substrates. The monkeys crouched bipedally in trees, but did not use other bipedal postures in trees. While on terrestrial substrates, they also crouched bipedally but occasionally stood upright and moved bipedally with orthograde posture. Bearded capuchin monkeys' behavior supports the suggestion from anatomical analysis that S. libidinosus is morphologically better adapted than its congeners to adopt orthograde postures.     

Sex differences and social organization in free-ranging spider monkeys (Ateles geoffroyi)

Several aspects of the social system of spider monkeys remain poorly understood in spite of previous studies of their behavior. Our work investigates sex differences of adultAteles geoffroyi to develop a better understanding of their social organization. A six-month field study of this species in Guatemala showed that adult males were both more aggressive and more socially cohesive than females, as well as more territorial. Adult females were more vocal, more submissive, more nonsocial, and more dispersed than adult males. Males were more likely to associate affinitively with other males than with females, and to direct their aggressive behaviors at females rather than males. Spider monkey society was found to be sex-segregated; males traveling and interacting in all-male subgroups, while females travel alone or with offspring. These findings are used, in conjunction with other evidence, to draw inferences about the dynamics of theAteles social system, and to derive an explanation for the evolution of spider monkey social organization. The frugivorous diet ofAteles is linked to the dispersion females and to the cohesion of related adult males, who form cooperative territorial groups, in which the low level of male-male competition is related to the absence of sexual dimorphism. Spider monkeys provide an illuminating contrast to the general primate model, derived from Old World monkeys, which links sexual dimorphism in size to sex differences in behavior, and ultimately to sexual selection.     

Habitat,diet, and activity patterns of free-ranging woolly spider monkeys (Brachyteles arachnoides E. Geoffroy 1806)

Results of a 10-month study of the ecology and behavior of free- ranging woolly spider monkeys (Brachyteles arachnoides)in Brazil show that these animals are strongly folivorous. Leaf-eating accounted for more than 50% of the total feeding time in all samples but one and accounted for more than 80% of the total feeding time in three samples. Mature foliage was routinely eaten. Woolly spider monkeys consistently spend more than 50% of each day quietly resting and sleeping. Animals travel little except when actively feeding and show low levels of social interaction. Such an activity profile suggests that woolly spider monkeys may often be living near the limits of their energetic resources. The social organization of the species is unusual for a folivorous primate in that small groups of females and associated immature animals confine their activities to discrete home-range areas, whereas males are itinerant, traveling over the home ranges of various female groups. Animals sharing a common home-range area show no permanent daily pattern of association other than that of mother-dependent offspring. Foraging alone or with few conspecifics should maximize each individual’s returns from foraging by minimizing the day range that must be traveled each day to locate foods while simultaneously lowering interference competition for higher-quality dietary resources.     

Seasonal Differences in Activity Patterns of Geoffroyi´s Spider Monkeys (<Emphasis Type="BoldItalic">Ateles geoffroyi</Emphasis>) Living in Continuous and Fragmented Forests in Southern Mexico

Understanding how primates adjust their behavior in response to seasonality in both continuous and fragmented forests is a fundamental challenge for primatologists and conservation biologists. During a 15-mo period, we studied the activity patterns of 6 communities of spider monkeys (Ateles geoffroyi) living in continuous and fragmented forests in the Lacandona rain forest, Mexico. We tested the effects of forest type (continuous and fragmented), season (dry and rainy), and their interaction on spider monkey activity patterns. Overall, monkeys spent more time feeding and less time traveling in fragments than in continuous forest. A more leafy diet and the spatial limitations in fragments likely explain these results. Time spent feeding was greater in the rainy than in the dry season, whereas time spent resting followed the opposite pattern. The increase in percent leaves consumed, and higher temperatures during the dry season, may contribute to the observed increase in resting time because monkeys probably need to reduce energy expenditure. Forest type and seasonality did not interact with activity patterns, indicating that the effect of seasonality on activities was similar across all sites. Our findings confirm that spider monkeys are able to adjust their activity patterns to deal with food scarcity in forest fragments and during the dry season. However, further studies are necessary to assess if these shifts are adequate to ensure their health, fitness, and long-term persistence in fragmented habitats.