Influences on the Feeding Behavior of Three Mammals in the Maruyama Zoo: Bears,Elephants, and Chimpanzees

Modifications of feeding conditions are essential to the establishment of environmental enrichment in zoos. In this study, we attempted to increase the duration of feeding by varying feeding conditions, such as the spatial or temporal distribution of foods, while keeping the sorts and amounts of food the same. Subjects included nonhuman animals reared at the Maruyama Zoo in Sapporo, Japan: 3 bears (Ursus arctos), 2 elephants (Elephas maximus), and 5 chimpanzees (Pan troglodytes). Considering that the feeding ecologies of these animals differed a great deal from those in the wild, we used the focal animal sampling method. Consisting of the spatial dispersive and massed feeding conditions, the experiment with bears found feeding time increased more in the dispersive condition. In addition, the behavioral rhythm of alternate feeding and sleeping appeared. The experiment with elephants, consisting of the temporal dispersive and massed condition, increased feeding time under the massed condition. The experiment using chimpanzees reversed conditions used in the elephant study and increased feeding time under the dispersive condition. Results indicated that these simple modifications both influenced an increasing duration of feeding and affected behavioral patterns in a day.     

Nutritional Consequences of Experimentally Introduced Tourism in Brown Bears

Abstract: Although numerous studies have documented behavioral effects of nature-based tourism on wildlife populations, few studies have determined whether behavioral changes translate to effects on individual condition and population health. This issue is currently a concern for wildlife managers in Alaska, USA, and Canada where bear viewing is a rapidly growing industry expanding into previously undisturbed bear habitats. Rather than record observations at long established tourism sites, we experimentally introduced bear viewing into 2 relatively undisturbed brown bear (Ursus arctos) populations in south-central Alaska. We examined the nutritional consequences of behavioral changes induced by the presence and activity of bear viewers for bears feeding on early summer vegetation and late-summer salmon (Oncorhynchus kisutch and O. nerka). We used Global Positioning System collars, monitored food resource availability, and quantified individual resource use and condition for a year prior to and during the introduction of bear viewing. Though bear viewing altered spatiotemporal resource use in all treatments, total resource use declined only when we exposed bears to 24-hour daily human activity. Energy expenditure, indexed as daily travel distances, was significantly higher when bears responded by altering spatial rather than temporal resource use. However, body weight and composition were unaffected by all treatments as bears shifted their foraging to other locations or times. Managers can minimize nutritional impacts of bear-viewing programs by avoiding spatial displacement and providing predictable time periods when bears can access food resources free of human activity. Bears in this study exhibited a high degree of behavioral plasticity, which may be an important factor in identifying flagship species for sustainable ecotourism programs.     

Exploitation of underground bee nests by three sympatric consumers in Loango National Park,Gabon

Honey represents a highly nutritious resource for animals, but is difficult to obtain given bees' defensive strategies. We investigated exploitation of the underground nests of stingless bees (Meliplebeia lendliana) by three sympatric consumers in Loango National Park, Gabon: the central African chimpanzee (Pan troglodytes troglodytes), forest elephant (Loxodonta cyclotis) and honey badger (Mellivora capensis). Given the differences in their respective morphological traits and sensory abilities, we hypothesized that chimpanzees would be more limited in digging out the bee nests, compared to the other two competitors, and would show behavioral strategies to overcome such constraints. Our dataset comprised camera trap footage recorded over 60 mo at 100 different bee nests. Chimpanzees visited the nests more often than the other consumers, showing a frequency of extraction success comparable to that observed in honey badgers, the most efficient digger. Both chimpanzees and honey badgers increased their extractive attempts across the dry season, whereas elephants did not. The soil hardness was greater during the dry season than the wet season and, possibly in order to compensate for this, chimpanzees showed a tendency toward digging at nests found in relatively softer soil. They also seemed to be inhibited by indirect cues left by other consumers, possibly as a risk‐avoidance strategy. Overall, chimpanzees and honey badgers extracted the underground nests of stingless bees with similar frequencies, whilst forest elephants did so only occasionally. Moreover, chimpanzees can use tools and other behavioral strategies to overcome the physical limitations that may constrain their exploitation of this resource.     

The sexually active states of free-ranging male African elephants (Loxodonta africana): defining musth and non-musth using endocrinology, physical signals, and behavior

Musth in male African elephants, Loxodonta africana, is associated with increased aggressive behavior, continuous discharge of urine, copious secretions from the swollen temporal glands, and elevated androgen levels. During musth, bulls actively seek out and are preferred by estrous females although sexual activity is not restricted to the musth condition. The present study combines recently established methods of fecal hormone analysis with long-term observations on male-female associations as well as the presence and intensity of physical signals to provide a more detailed picture about the physical, physiological, and behavioral characteristics of different states of sexual activity in free-ranging African elephants. Based on quantitative shifts in individual bull association patterns, the presence of different physical signals, and significant differences in androgen levels, a total of three potential sub-categories for sexually active bulls could be established. The results demonstrate that elevations in androgen levels are only observed in sexually active animals showing temporal gland secretion and/or urine dribbling, but are not related to the age of the individual. Further, none of the sexually active states showed elevated glucocorticoid output indicating that musth does not represent an HPA-mediated stress condition. On the basis of these results, we suggest that the term "musth" should be exclusively used for the competitive state in sexually active male elephants and that the presence of urine dribbling should be the physical signal used for defining this state.     

Brief communication: arboreal bipedalism in Bwindi chimpanzees

Evidence of the form and function of bipedal behavior in nonhuman primates provides critical evidence to test theories about the origins of hominid bipedalism. Bipedalism has long been considered an evolutionarily interesting but rare behavior in wild chimpanzees. During May 2001, chimpanzees of the Ruhija community in the Bwindi Impenetrable National Park, Uganda, engaged in an exceptional frequency of arboreal bipedalism when feeding in large Ficus trees. Seventy-eight bipedal bouts of at least 5 sec duration were recorded for the entire community (0.49 bouts/hr), with a mean duration of 13.7 sec (+/-1.6 sec). The animals employed many variations on the bipedal postural theme, ranging from erect standing on the largest substrates while grasping overhead limbs for support, to standing on one leg while suspending the other leg in space, to extended-lean standing, in which bipedal standing transitioned into horizontal arm-leg suspension as the animal reached for more distant fruits. Bipedalism was used as part of a behavioral repertoire that integrated brachiation, four-limbed suspension, and forelimb-supported standing for effective small-fruit foraging. These observations suggest that under certain ecological conditions, arboreal bipedalism can be an important posture for wild chimpanzees, and may have been an important behavioral precursor to full terrestrial bipedalism.     

A preliminary study of the behavioral effects of feeding enrichment on African elephants

Although commonly practiced with many species, feeding enrichment in elephants is understudied. The behavioral effects of feeding enrichment were tested in three African elephants by substituting an equal dry weight of browse for hay in an ABA design. The results showed a significant increase in feeding and significant decreases in drinking and inactivity when the browse was present. Additionally, changes in feeding, inactivity, and time spent in contact were observed outside the time the browse was actually presented, although the relationship of these changes to the experimental methodology is unknown. A significant increase in visibility to zoo visitors during the browse conditions demonstrates that browse is an effective, naturalistic method for increasing visibility as well as species-typical behaviors. Managers of captive elephants should consult with a nutritionist to address issues of energetics, nutritional content, and secondary compounds when using browse as feeding enrichment. Zoo Biol 19:485-493, 2000. Copyright 2000 Wiley-Liss, Inc.     

Physical, physiological, and behavioral correlates of musth in captive African elephants (Loxodonta africana)

Although musth in male African elephants (Loxodonta africana) is known to be associated with increased aggressiveness, urine dribbling (UD), temporal gland secretion (TGS), and elevated androgens, the temporal relationship between these changes has not been examined. Here, we describe the pattern of musth-related characteristics in 14 captive elephant bulls by combining long-term observations of physical and behavioral changes with physiological data on testicular and adrenal function. The length of musth periods was highly variable but according to our data set not related to age. Our data also confirm that musth is associated with elevated androgens and, in this respect, show that TGS and UD are downstream effects of this elevation, with TGS responding earlier and to lower androgen levels than UD. Because the majority of musth periods were associated with a decrease in glucocorticoid levels, our data also indicate that musth does not represent a physiological stress mediated by the hypothalamic-pituitary-adrenal axis. Furthermore, we demonstrate that the occurrence of musth is associated with increased aggression and that this is presumably androgen mediated because aggressive males had higher androgen levels. Collectively, the information generated contributes to a better understanding of what characterizes and initiates musth in captive African elephants and provides a basis for further studies designed to examine in more detail the factors regulating the intensity and duration of musth.     

Testing for personality consistency across naturally occurring behavioral contexts in sanctuary chimpanzees (Pan troglodytes)

Personality is both a reflection of the bio-behavioral profile of individuals and a summary of how they typically interact with their physical and social world. Personality is usually defined as having distinct behavioral characteristics, which are assumed to be consistent over time and across contexts. Like other mammals, primates have individual differences in personality. Although temporal consistency is sometimes measured in primates, and contextual consistency is sometimes measured across experimental contexts, it is rare to measure both in the same individuals and outside of experimental settings. Here, we aim to measure both temporal and contextual consistency in chimpanzees, assessing their personality with behavioral observations from naturally occurring contexts (i.e., real-life settings). We measured personality-based behaviors in 22 sanctuary chimpanzees, in the contexts of feeding, affiliation, resting, and solitude, across two time periods, spanning 4 years. Of the 22 behaviors recorded, about 64% were consistent across two to four contexts and 50% were consistent over time. Ten behaviors loaded significantly onto three trait components: explorativeness, boldness-sociability, and anxiety-sociability, as revealed by factor analysis. Like others, we documented individual differences in the personality of chimpanzees based on reliably measured observations in real-life contexts. Furthermore, we demonstrated relatively strong, but not absolute, temporal, and contextual consistency in personality-based behaviors. We also found another aspect of individual differences in personality, specifically, the extent to which individual chimpanzees show consistency. Some individuals showed contextual and temporal consistency, whereas others show significant variation across behaviors, contexts, and/or time. We speculate that the relative degree of consistency in personality may vary within chimpanzees. It may be that different primate species vary in the extent to which individuals show consistency of personality traits. Our behavioral-based assessment can be used with wild populations, increasing the validity of personality studies, facilitating comparative studies and potentially being applicable to conservation efforts.     

Urinary C-peptide tracks seasonal and individual variation in energy balance in wild chimpanzees

C-peptide of insulin presents a promising new tool for behavioral ecologists that allows for regular, non-invasive assessment of energetic condition in wild animals. C-peptide is produced on an equimolar basis with insulin, thus is indicative of the body's response to available glucose and, with repeated measurement, provides a biomarker of energy balance. As yet, few studies have validated the efficacy of C-peptide for monitoring energy balance in wild animals. Here, we assess seasonal and interindividual variation in urinary C-peptide concentrations of East African chimpanzees (Pan troglodytes schweinfurthii). We assayed 519 urine samples from 13 adult male chimpanzees in the Kanyawara community of Kibale National Park, Uganda. C-peptide levels were significantly predicted by the total amount of fruit and the amount of preferred fruit in the diet. However, chimpanzees had very low C-peptide titers during an epidemic of severe respiratory illness, despite highly favorable feeding conditions. Kanyawara males had significantly lower C-peptide levels than males at Ngogo, a nearby chimpanzee community occupying a more productive habitat. Among Kanyawara males, low-ranking males had consistently higher C-peptide levels than dominant males. While counterintuitive, this result supports previous findings of costs associated with dominance in male chimpanzees. Our preliminary investigations demonstrate that C-peptide has wide applications in field research, providing an accessible tool for evaluating seasonal and individual variation in energetic condition, as well as the costs of processes such as immune function and reproduction.     

Feeding Consequences of Hand and Foot Disability in Wild Adult Chimpanzees (<Emphasis Type="Italic">Pan troglodytes schweinfurthii</Emphasis>)

Many wild primates experience long-term limb disability, and their ability to cope with disability has implications for survival and fitness. We quantified the arboreal feeding and postural behaviors of adult chimpanzees to study the consequences of physical limb disabilities. We collected behavioral data for a total of 8 mo on chimpanzees at Sebitoli, Kibale National Park, Uganda, focusing on the time spent feeding, common feeding tree species, body postures, and substrate use. Of the 51 chimpanzees we observed, 16 (31%) exhibited limb anomalies, which varied in form and severity. Disabled chimpanzees climbed as high as chimpanzees without disabilities and did not differ from nondisabled chimpanzees in the amount they used feeding tree species. Adult chimpanzees with severe hand disability spent significantly more time feeding than nondisabled individuals. In addition, manually disabled adults did not suspend themselves from branches during feeding as frequently as nondisabled adults and used larger substrates for gripping and sitting than nondisabled adults. These results indicate that disabled individuals compensate to carry out feeding activities in trees.     

Habitat‐mediated timing of migration in polar bears: an individual perspective

Migration phenology is largely determined by how animals respond to seasonal changes in environmental conditions. Our perception of the relationship between migratory behavior and environmental cues can vary depending on the spatial scale at which these interactions are measured. Understanding the behavioral mechanisms behind population‐scale movements requires knowledge of how individuals respond to local cues. We show how time‐to‐event models can be used to predict what factors are associated with the timing of an individual's migratory behavior using data from GPS collared polar bears (Ursus maritimus) that move seasonally between sea ice and terrestrial habitats. We found the concentration of sea ice that bears experience at a local level, along with the duration of exposure to these conditions, was most associated with individual migration timing. Our results corroborate studies that assume thresholds of >50% sea ice concentration are necessary for suitable polar bear habitat; however, continued periods (e.g., days to weeks) of exposure to suboptimal ice concentrations during seasonal melting were required before the proportion of bears migrating to land increased substantially. Time‐to‐event models are advantageous for examining individual movement patterns because they account for the idea that animals make decisions based on an accumulation of knowledge from the landscapes they move through and not simply the environment they are exposed to at the time of a decision. Understanding the migration behavior of polar bears moving between terrestrial and marine habitat, at multiple spatiotemporal scales, will be a major aspect of quantifying observed and potential demographic responses to climate‐induced environmental changes.     

Influence of food distribution on behavior in captive bongos,Taurotragus euryceros: An experimental investigation

A feeding experiment was conducted to test if Bongo antelopes, being selective feeders searching for clumped quality food, show similar behavior changes as primates under similar conditions. One group of 3 females and 3 calves of Dvur Kralove Zoo was either fed clumped (all food, i.e., leaves and branches of 4 species of tree plus rye grass in one heap), or the same amount distributed over as many heaps as animals present. No increase in agonistic interactions under clumped conditions was found, but non-agonistic and sociopositive patterns increased significantly. Intake of low quality food increased under clumped conditions. Social distances and enclosure use were not significantly different. These findings are in accordance with predictions from models developed for primate feeding strategies, and suggest that selectively feeding antelopes possess behavioral mechanisms for tension-reduction similar to “contest-type” primates. Zoo Biol 16:237–245, 1997. © 1997 Wiley-Liss, Inc.     

Physiological stress and refuge behavior by African elephants

Physiological stress responses allow individuals to adapt to changes in their status or surroundings, but chronic exposure to stressors could have detrimental effects. Increased stress hormone secretion leads to short-term escape behavior; however, no studies have assessed the potential of longer-term escape behavior, when individuals are in a chronic physiological state. Such refuge behavior is likely to take two forms, where an individual or population restricts its space use patterns spatially (spatial refuge hypothesis), or alters its use of space temporally (temporal refuge hypothesis). We tested the spatial and temporal refuge hypotheses by comparing space use patterns among three African elephant populations maintaining different fecal glucocorticoid metabolite (FGM) concentrations. In support of the spatial refuge hypothesis, the elephant population that maintained elevated FGM concentrations (iSimangaliso) used 20% less of its reserve than did an elephant population with lower FGM concentrations (Pilanesberg) in a reserve of similar size, and 43% less than elephants in the smaller Phinda reserve. We found mixed support for the temporal refuge hypothesis; home range sizes in the iSimangaliso population did not differ by day compared to nighttime, but elephants used areas within their home ranges differently between day and night. Elephants in all three reserves generally selected forest and woodland habitats over grasslands, but elephants in iSimangaliso selected exotic forest plantations over native habitat types. Our findings suggest that chronic stress is associated with restricted space use and altered habitat preferences that resemble a facultative refuge behavioral response. Elephants can maintain elevated FGM levels for ≥ 6 years following translocation, during which they exhibit refuge behavior that is likely a result of human disturbance and habitat conditions. Wildlife managers planning to translocate animals, or to initiate other management activities that could result in chronic stress responses, should consider the potential for, and consequences of, refuge behavior.     

Structure and Dynamics of Minke Whale Surfacing Patterns in the Gulf of St. Lawrence,Canada

Animal behavioral patterns can help us understand physiological and ecological constraints on animals and its influence on fitness. The surfacing patterns of aquatic air-breathing mammals constitute a behavioral pattern that has evolved as a trade-off between the need to replenish oxygen stores at the surface and the need to conduct other activities underwater. This study aims to better understand the surfacing pattern of a marine top predator, the minke whale (Balaenoptera acutorostrata), by investigating how their dive duration and surfacing pattern changes across their activity range. Activities were classified into resting, traveling, surface feeding and foraging at depth. For each activity, we classified dives into short and long dives and then estimated the temporal dependence between dive types. We found that minke whales modified their surfacing pattern in an activity-specific manner, both by changing the expression of their dives (i.e. density distribution) and the temporal dependence (transition probability) between dive types. As the depth of the prey layer increased between activities, the surfacing pattern of foraging whales became increasingly structured, going from a pattern dominated by long dives, when feeding at the surface, to a pattern where isolated long dives were followed by an increasing number of breaths (i.e. short dives), when the whale was foraging at depth. A similar shift in surfacing pattern occurred when prey handling time (inferred from surface corralling maneuvers) increased for surface feeding whales. The surfacing pattern also differed between feeding and non-feeding whales. Resting whales did not structure their surfacing pattern, while traveling whales did, possibly as a way to minimize cost of transport. Our results also suggest that minke whales might balance their oxygen level over multiple, rather than single, dive cycles.     

Visual Cues Given by Humans Are Not Sufficient for Asian Elephants (Elephas maximus) to Find Hidden Food

Recent research suggests that domesticated species – due to artificial selection by humans for specific, preferred behavioral traits – are better than wild animals at responding to visual cues given by humans about the location of hidden food. \Although this seems to be supported by studies on a range of domesticated (including dogs, goats and horses) and wild (including wolves and chimpanzees) animals, there is also evidence that exposure to humans positively influences the ability of both wild and domesticated animals to follow these same cues. Here, we test the performance of Asian elephants (Elephas maximus) on an object choice task that provides them with visual-only cues given by humans about the location of hidden food. Captive elephants are interesting candidates for investigating how both domestication and human exposure may impact cue-following as they represent a non-domesticated species with almost constant human interaction. As a group, the elephants (n = 7) in our study were unable to follow pointing, body orientation or a combination of both as honest signals of food location. They were, however, able to follow vocal commands with which they were already familiar in a novel context, suggesting the elephants are able to follow cues if they are sufficiently salient. Although the elephants’ inability to follow the visual cues provides partial support for the domestication hypothesis, an alternative explanation is that elephants may rely more heavily on other sensory modalities, specifically olfaction and audition. Further research will be needed to rule out this alternative explanation.     

Behavioral plasticity mitigates the effect of warming on white‐tailed deer

Climate change is expected to create novel environments in which extant species cannot persist, therefore leading to the loss of them and their associated ecological functions within the ecosystem. However, animals may employ behavioral mechanisms in response to warming that could allow them to maintain their functional roles in an ecosystem despite changed temperatures. Specifically, animals may shift their activity in space or time to make use of thermal heterogeneity on the landscape. However, few studies consider the role of behavioral plasticity and spatial or temporal heterogeneity in mitigating the effects of climate change. We conducted experiments to evaluate the potential importance of behavior in mediating the net effects of warming on white‐tailed deer (Odocoileus virginianus). We used shade structures to manipulate the thermal environment around feeding stations to monitor deer feeding activity and measure total consumption. In individual experiments where deer only had access to unshaded feeders, deer fed less during the day but compensated by increasing feeding during times when temperature was lower. In group experiments where deer had access to both shaded and unshaded feeders, deer often fed during the day but disproportionally preferred the cooler, shaded feeders. Our results suggest that deer can capitalize on temporal and spatial heterogeneity in the thermal environment to meet nutritional and thermal requirements, demonstrating the importance of behavioral plasticity when predicting the net effects of climate change.     

The daily activity of the elephant in the Rwenzori National Park, Uganda

Eighteen elephants were followed individually in the Rwenzori National Park, Uganda for varying periods and their activity noted at 4-min intervals. Continuous data extending over 24 h or more were collected from seven of these elephants. About 75% of the total time was spent in feeding and about 75% of the feeding activity consisted of grazing or feeding at ground level. There were three feeding peaks, one in the morning, another in the afternoon and the third around midnight. The feeding rate tended to increase during the day. The proportion of the total feeding time spent in browsing increased in the dry season. Walking took place mainly at dusk. The principal sleeping period occurred during the small hours of the morning with a shorter rest period in the early afternoon. The elephants defaecated about eleven times in the 24 h and drank an average of 1–3 times. The rate of progress through the bush averaged 0–5 km/h. Comparisons are made with published data concerning the activity patterns of waterbuck and warthog in the Rwenzori Park and of elephants in the Tsavo National Park, Kenya.     

Collared peccary (Pecari tajacu) behavioral reactions toward a dead member of the herd

Humans, elephants, chimpanzees, and cetaceans show concern with the death of other members of their species and respond to death in particular ways. Science considers that these species are exceptions and that other mammal species show little or no reaction to the dead bodies of individuals of their species. Collared peccaries (Pecari tajacu; Tayassuidae) are social animals that live in groups of 5–50 individuals maintaining close and complex social relationships. The collared peccary occupies many different environments and it is widely distributed from the south of North America to the north of Argentina. Their behavior is well studied, but we know little about their behavior toward the dead. We directly observed and filmed with a camera trap the reactions of a five‐member herd of collared peccaries to the death of a herd member. We worked on a suburban forested area in the mountains of central Arizona. We found that the herd visited and spent time with the dead body for 10 days after the peccary died. The frequency of the visits declined until the cadaver was consumed by coyotes. Most of the videos showed two individuals visited the dead animal (44%), solitary records were also frequent (39%) and only 4% of the videos recorded three peccaries. Visits were more frequent during the night (64%). Peccaries do react to the death of a herd member by behaving in particular ways. Reactions include pushing at the dead individual, staring at it, biting it, and trying to pick it up by putting their snout under the corpse and pushing it up, and defending it from coyotes, among others. These levels of behavioral complexity for peccaries are beyond those known so far. The behaviors of this herd of peccaries resemble those of humans, cetaceans, chimpanzees, and elephants and show that these groups are not the only ones that react to death.     

Effective sociodemographic population assessment of elusive species in ecology and conservation management

Wildlife managers are urgently searching for improved sociodemographic population assessment methods to evaluate the effectiveness of implemented conservation activities. These need to be inexpensive, appropriate for a wide spectrum of species and straightforward to apply by local staff members with minimal training. Furthermore, conservation management would benefit from single approaches which cover many aspects of population assessment beyond only density estimates, to include for instance social and demographic structure, movement patterns, or species interactions. Remote camera traps have traditionally been used to measure species richness. Currently, there is a rapid move toward using remote camera trapping in density estimation, community ecology, and conservation management. Here, we demonstrate such comprehensive population assessment by linking remote video trapping, spatially explicit capture–recapture (SECR) techniques, and other methods. We apply it to three species: chimpanzees Pan troglodytes troglodytes, gorillas Gorilla gorilla gorilla, and forest elephants Loxodonta cyclotis in Loango National Park, Gabon. All three species exhibited considerable heterogeneity in capture probability at the sex or group level and density was estimated at 1.72, 1.2, and 1.37 individuals per km² and male to female sex ratios were 1:2.1, 1:3.2, and 1:2 for chimpanzees, gorillas, and elephants, respectively. Association patterns revealed four, eight, and 18 independent social groups of chimpanzees, gorillas, and elephants, respectively: key information for both conservation management and studies on the species' ecology. Additionally, there was evidence of resident and nonresident elephants within the study area and intersexual variation in home range size among elephants but not chimpanzees. Our study highlights the potential of combining camera trapping and SECR methods in conducting detailed population assessments that go far beyond documenting species diversity patterns or estimating single species population size. Our study design is widely applicable to other species and spatial scales, and moderately trained staff members can collect and process the required data. Furthermore, assessments using the same method can be extended to include several other ecological, behavioral, and demographic aspects: fission and fusion dynamics and intergroup transfers, birth and mortality rates, species interactions, and ranging patterns.     

Behavioral seasonality in Mahale chimpanzees

To analyze how the chimpanzees in the Mahale Mountains National Park, Tanzania, change their grouping pattern, activity budget, travel speed, and travel distance within an annual cycle, I divided 1-year data into four periods. The Mahale chimpanzees have the behavioral flexibility to adapt to various climates and exhibited at least three behavioral seasons. In the early wet season, chimpanzees formed a few, large parties, and spent much time feeding on insects and animal meat. In the early and late dry seasons, chimpanzees maintained party sizes as large as in the early wet season, and traveled distances similar to the early wet season, but spent the most time feeding and traveling within the year. By contrast, in the late wet season chimpanzee parties broke up into more numerous, small groups, and traveled slowly over shorter distances. Although time spent feeding and traveling were the same as that in the early wet season, time spent feeding on terrestrial herbaceous vegetation (THV) was the highest in the year. The results suggest that chimpanzees travel longer, faster, and farther in seasons when they form large parties.