Comparison of instantaneous and locomotor bout sampling methods: a case study of adult male chimpanzee locomotor behavior and substrate use.

Currently two methods, instantaneous and locomotor bout sampling, are used most commonly in studies of locomotor behavior. To date, no study has addressed how comparable the results of the two methods are. This paper considers whether different sampling methods of locomotor behavior produce different results. Continuous locomotor bout and instantaneous sampling were carried out simultaneously on each focal animal during a seven month study of chimpanzee positional behavior in the Tai Forest of the Ivory Coast. Results provide two independent sets of data which describe the same events. Results indicate that as locomotor bouts are frequently presented (the percentage of bouts spent in an activity), they overrepresent the frequencies of activities that occur relatively often, but for short distances, and underrepresent activities that have a relatively large mean distance per bout. However, when bouts are weighted with distance, as originally defined by Fleagle (1976b), there are no significant differences in the results obtained by the two methods. Both provide similar results for the frequencies of locomotor activities, frequency of substrate use, and the relationship between substrate and locomotor activity. The advantage of instantaneous sampling is that because it is done at regular intervals, it can easily be carried out in conjunction with sampling other behavioral and ecological data. The advantages of locomotor bout sampling are that it permits the sampling of rare or brief locomotor events and allows for an analysis of sequences of locomotor activities. This paper demonstrates that the two methods can be conducted simultaneously and thus provide the richest return from which the effect of environment and morphology on locomotion can be addressed.     

A survey of entodiniomorphid ciliates in chimpanzees and bonobos

Intestinal entodiniomorphid ciliates are commonly diagnosed in the feces of wild apes of the genera Pan and Gorilla. Although some authors previously considered entodiniomorphid ciliates as possible pathogens, a symbiotic function within the intestinal ecosystem and their participation in fiber fermentation has been proposed. Previous studies have suggested that these ciliates gradually disappear under captive conditions. We studied entodiniomorphid ciliates in 23 captive groups of chimpanzees, three groups of captive bonobos and six populations of wild chimpanzees. Fecal samples were examined using Sheather's flotation and Merthiolate‐Iodine‐Formaldehyde Concentration (MIFC) methods. We quantified the number of ciliates per gram of feces. The MIFC method was more sensitive for ciliate detection than the flotation method. Ciliates of genus Troglodytella were detected in 13 groups of captive chimpanzees, two groups of bonobos and in all wild chimpanzee populations studied. The absence of entodiniomorphids in some captive groups might be because of the extensive administration of chemotherapeutics in the past or a side‐effect of the causative or prophylactic administration of antiparasitic or antibiotic drugs. The infection intensities of ciliates in captive chimpanzees were higher than in wild ones. We suppose that the over‐supply of starch, typical in captive primate diets, might induce an increase in the number of ciliates. In vitro studies on metabolism and biochemical activities of entodiniomorphids are needed to clarify their role in ape digestion. Am J Phys Anthropol 2010. © 2009 Wiley‐Liss, Inc.     

Tolerant food sharing and reciprocity is precluded by despotism among bonobos but not chimpanzees

Tolerant food sharing among human foragers can largely be explained by reciprocity. In contrast, food sharing among chimpanzees and bonobos may not always reflect reciprocity, which could be explained by different dominance styles: in egalitarian societies reciprocity is expressed freely, while in more despotic groups dominants may hinder reciprocity. We tested the degree of reciprocity and the influence of dominance on food sharing among chimpanzees and bonobos in two captive groups. First, we found that chimpanzees shared more frequently, more tolerantly, and more actively than bonobos. Second, among chimpanzees, food received was the best predictor of food shared, indicating reciprocal exchange, whereas among bonobos transfers were mostly unidirectional. Third, chimpanzees had a shallower and less linear dominance hierarchy, indicating that they were less despotic than bonobos. This suggests that the tolerant and reciprocal sharing found in chimpanzees, but not bonobos, was made possible by the absence of despotism. To investigate this further, we tested the relationship between despotism and reciprocity in grooming using data from an additional five groups and five different study periods on the main groups. The results showed that i) all chimpanzee groups were less despotic and groomed more reciprocally than bonobo groups, and ii) there was a general negative correlation between despotism and grooming reciprocity across species. This indicates that an egalitarian hierarchy may be more common in chimpanzees, at least in captivity, thus fostering reciprocal exchange. We conclude that a shallow dominance hierarchy was a necessary precondition for the evolution of human‐like reciprocal food sharing. Am J Phys Anthropol 143:41–51, 2010. © 2010 Wiley‐Liss, Inc.     

Sex differences in adult chimpanzee positional behavior: The influence of body size on locomotion and posture

Focal animal instantaneous sampling of adult male and female chimpanzee positional behavior was conducted during a 7-month study in the Tai Forest, Ivory Coast, in order to determine whether there are sex differences in the locomotion, posture, substrate use, and height preference of sexually dimorphic adult chimpanzees, and if so, whether these differences support predictions based on body size differences. Results indicate that as predicted, adult male and female chimpanzees differ in their arboreal locomotor behavior, with the larger males using less quadrupedalism and more climbing, scrambling, and aided bipedalism than females during feeding locomotion. There is a sex difference in height preference as well, with female chimpanzees consistently using more arboreal behavior than males, primarily during resting. Although it has been previously demonstrated that separate primate species of differing body size differ in locomotor and postural activities (Fleagle and Mittermeier, 1980; Crompton, 1984), this study clearly demonstrates that body size differences within a species can also be correlated with differences in locomotor behavior. These findings may influence how we interpret sex differences in body size of extinct species. © 1993 Wiley-Liss, Inc.     

Brief Communication: Quantitative‐ and molecular‐genetic differentiation in humans and chimpanzees: Implications for the evolutionary processes underlying cranial diversification

Estimates of the amount of genetic differentiation in humans among major geographic regions (e.g., Eastern Asia vs. Europe) from quantitative‐genetic analyses of cranial measurements closely match those from classical‐ and molecular‐genetic markers. Typically, among‐region differences account for ～10% of the total variation. This correspondence is generally interpreted as evidence for the importance of neutral evolutionary processes (e.g., genetic drift) in generating among‐region differences in human cranial form, but it was initially surprising because human cranial diversity was frequently assumed to show a strong signature of natural selection. Is the human degree of similarity of cranial and DNA‐sequence estimates of among‐region genetic differentiation unusual? How do comparisons with other taxa illuminate the evolutionary processes underlying cranial diversification? Chimpanzees provide a useful starting point for placing the human results in a broader comparative context, because common chimpanzees (Pan troglodytes) and bonobos (Pan paniscus) are the extant species most closely related to humans. To address these questions, I used 27 cranial measurements collected on a sample of 861 humans and 263 chimpanzees to estimate the amount of genetic differentiation between pairs of groups (between regions for humans and between species or subspecies for chimpanzees). Consistent with previous results, the human cranial estimates are quite similar to published DNA‐sequence estimates. In contrast, the chimpanzee cranial estimates are much smaller than published DNA‐sequence estimates. It appears that cranial differentiation has been limited in chimpanzees relative to humans. Am J Phys Anthropol 154:615–620, 2014. © 2014 Wiley Periodicals, Inc.     

Inhibition of social behavior in chimpanzees under high-density conditions

This is the first study to investigate the short-term effects of high population density on captive chimpanzees (Pan troglodytes). Subjects of the study were 45 chimpanzees living in five different groups at the Yerkes Regional Primate Research Center. The groups were observed under two conditions: 1) when they had access to both the indoor and outdoor sections of their enclosures; 2) during cold days when they were locked into the indoor runs, which reduced the available space by more than half. Under the high-density condition, allogrooming and submissive greetings decreased, but juvenile play increased. Remarkably, the rate of various forms of agonistic behavior, such as aggression, bluff charge, bluff display, and hooting, occurred less frequently under the high-density condition. This general decrease in adult social activity, including agonistic behavior, can be interpreted as an inhibition strategy to reduce opportunities for conflict when interindividual distances are reduced. This strategy is probably effective only in the short run, however. Behavioral indicators of anxiety, such as rough scratching and yawning, showed elevated rates, suggesting increased social tension under the high-density condition. Am. J. Primatol. 41:213–228, 1997. © 1997 Wiley-Liss, Inc.     

Locomotor behavior of Pan paniscus in Zaire

The locomotor behavior of Pan paniscus was studied over a four-week period in Equateur, Republic of Zaire. Bonobos were found to be both arboreal and terrestrial in their daily activities. In the trees adult bonobos are basically quadrupedal, but they also have significant components of armswinging, diving, leaping, and bipedalism in their locomotor repertoire.     

Energetic costs of territorial boundary patrols by wild chimpanzees

Chimpanzees are well known for their territorial behavior. Males who belong to the same community routinely patrol their territories, occasionally making deep incursions into those of their neighbors. Male chimpanzees may obtain several fitness benefits by participating in territorial boundary patrols, but patrolling is also likely to involve fitness costs. Patrollers risk injury or even death, and patrols may be energetically costly and may involve opportunity costs. Although territorial patrols have been reported at all long‐term chimpanzee study sites, quantitative data on their energetic costs have not previously been available. I evaluated the energy costs of patrolling for male chimpanzees at Ngogo, Kibale National Park, Uganda during 14 months of observation. In 29 patrols and matched control periods, I recorded the distances covered and time spent traveling and feeding by chimpanzees. I found that male chimpanzees covered longer distances, spent more time traveling, and spent less time feeding during patrols than during control periods. These results support the hypothesis that chimpanzees incur energetic costs while patrolling and suggest that ecological factors may constrain the ability of chimpanzees to patrol. Am. J. Primatol. 72:93–103, 2010. © 2009 Wiley‐Liss, Inc.     

Resocialization of a group of ex-laboratory chimpanzees, Pan troglodytes.

In order to determine if socially deprived chimpanzees could establish an integrated social unit and reproduce, a group of wild-born individuals from a single-cage laboratory environment were released onto a small island in a South Florida tourist park in 1975. The present study was undertaken during June and July of 1988 to evaluate the long term results of the experiment and to document the reproductive history of the chimpanzees since their release on the island. The evidence presented in this paper suggests that it is possible for laboratory chimpanzees to be resocialized and to reproduce successfully.     

Demographic influences on the hunting behavior of chimpanzees

We investigated hunting in an unusually large community of wild chimpanzees at Ngogo in the Kibale National Park, Uganda. Aspects of predation were recorded with respect to the prey, the predators, and hunting episodes. During 23 months of observation, the Ngogo chimpanzees caught 128 prey items from four primate and three ungulate species. Chimpanzees preyed selectively on immature red colobus primarily during group hunts, with adult males making the majority of kills. Party size and composition were significant predictors of the probability that chimpanzees would hunt and of their success during attempts. Chimpanzees were more likely to hunt red colobus if party size and the number of male hunters were large; party size and the number of male hunters were also significantly larger in successful compared with unsuccessful hunts. The Ngogo chimpanzees did not appear to hunt cooperatively, but reciprocal meat-sharing typically took place after kills. Hunts occurred throughout the year, though there was some seasonality as displayed by periodic hunting binges. The extremely high success rate and large number of kills made per successful hunt are the two most striking aspects of predation by the Ngogo chimpanzees. We compare currently available observations of chimpanzee hunting behavior across study sites and conclude that the large size of the Ngogo community contributes to their extraordinary hunting success. Demographic differences between groups are likely to contribute to other patterns of interpopulation variation in chimpanzee predation. Am J Phys Anthropol 109:439–454, 1999. © 1999 Wiley-Liss, Inc.     

Locomotor behavior and long bone morphology in individual free-ranging chimpanzees

We combine structural limb data and behavioral data for free-ranging chimpanzees from Ta? (Ivory Coast) and Mahale National Parks (Tanzania) to begin to consider the relationship between individual variation in locomotor activity and morphology. Femoral and humeral cross sections of ten individuals were acquired via computed tomography. Locomotor profiles of seven individuals were constructed from 3387 instantaneous time-point observations (87.4 hours). Within the limited number of suitable chimpanzees, individual variation in locomotor profiles displayed neither clear nor consistent trends with diaphyseal cross-sectional shapes. The percentages of specific locomotor modes did not relate well to diaphyseal shapes since neither infrequent nor frequent locomotor modes varied consistently with shapes. The percentage of arboreal locomotion, rather than estimated body mass, apparently had comparatively greater biological relevance to variation in diaphyseal shape. The mechanical consequences of locomotor modes on femoral and humeral diaphyseal shapes (e.g., orientation of bending strains) may overlap between naturalistic modes more than currently is recognized. Alternatively, diaphyseal shape may be unresponsive to mechanical demands of these specific locomotor modes. More data are needed in order to discern between these possibilities. Increasing the sample to include additional free-ranging chimpanzees, or primates in general, as well as devoting more attention to the mechanics of a greater variety of naturalistic locomotor modes would be fruitful to understanding the behavioral basis of diaphyseal shapes.     

Use of leaves as cushions to sit on wet ground by wild chimpanzees

A new type of tool use, leaf cushion, by wild chimpanzees (Pan troglodytes verus) at Bossou, Guinea, was found. We report two cases: one is indirect evidence; the other is direct observation of a chimpanzee who used the tool. Both cases indicate that chimpanzees used a set of leaves as a cushion while sitting on wet ground. Chimpanzees at Bossou show various kinds of tool use, some of which are unique to the community. Most of these behavioral patterns are subsistence tool use for obtaining food, as at other study sites. The use of leaves as a cushion adds to the few instances of nonsubsistence, elementary technology seen used by wild chimpanzees. Am. J. Primatol. 44:215–220, 1998. © 1998 Wiley-Liss, Inc.     

Utilization of space by adult and juvenile groups of captive chimpanzees (Pan troglodytes)

A systematic analysis of space utilization in the home enclosures of four adult and three juvenile social groups of chimpanzees (Pan troglodytes) was conducted at the Primate Foundation of Arizona. The locations of all members of each group (excluding nursing infants) were recorded several times per week over a 3-month period. Adult chimpanzees were observed significantly more often in small vs large cages, on upper vs lower levels, and near cage perimeters vs cage centers. Analyses of sex differences in adults revealed that adult females are largely responsible for these space use patterns; adult males show only a significantly higher usage of perimeter areas. Individual differences were found in the habitual use of certain enclosure sites. Juveniles show no spatial patterns with regard to the above parameters but appear to evenly use all available space. Both adults and juveniles were observed to be in contact with mesh barriers more often than with solid walls. These findings suggest important elements that should be incorporated into the caging design of captive chimpanzee facilities.     

The neighbor effect: Other groups influence intragroup agonistic behavior in captive chimpanzees

While intercommunity interactions are well documented in wild chimpanzees, the influence of neighboring captive groups on social behavior has not been investigated. This study examined the influence of vocalizations and noisy displays produced in neighboring groups (i.e., “neighbor vocalization”) on intragroup hooting, bluff displays, and agonistic behavior. Chimpanzees from two broadly differing housing conditions were selected in order to examine the effect of neighboring groups: “Wing Groups” (those living in indoor/outdoor pens with auditory access to large number of neighboring individuals) and “Building Groups” (those housed in one indoor building, able to hear the calls and noisy displays produced by a small number of neighboring individuals occupying the building). Data were collected on 58 adult and juvenile subjects living in social groups varying in size from two to 14 individuals. Observation sessions were divided into two categories depending on the level of neighbor vocalization. Wing Groups showed higher rates of hooting, bluff displays, and agonistic behavior against other group members when levels of neighbor vocalization were high. Building Groups showed the same pattern, with results for bluff displays reaching statistical significance. This effect was also found through an analysis of the time periods immediately surrounding incidents of neighbor vocalization: Hooting and bluff displays were significantly more common after than before neighbor vocalizations. In spite of variation between the two housing conditions with respect to overall levels of neighbor vocalization, degree of differences between the high and low vocalization levels, and methods of data collection, a consistent neighbor effect was detected. This effect is congruous with behavioral patterns observed in the wild, and is therefore considered an expression of species-typical behavior as opposed to an artifact of captivity. These results are relevant to captive management, and bear on the influence of housing condition on psychological well-being. © 1996 Wiley-Liss, Inc.     

The relationship between socio‐sexual behavior and salivary cortisol in bonobos: tests of the tension regulation hypothesis

Bonobos are known for their pacifistic behavior and their large repertoire of behaviors that are thought to serve conflict resolution. One is an unusual form of ventro‐ventral mounting that facilitates genital contacts (GC). Various hypotheses have been proposed to explain its function. In this study we tested predictions of the tension regulation hypothesis using salivary cortisol as a marker for social stress. The results indicate a temporal relationship between GC and cortisol levels. Compared with baseline data and matched samples of unrestricted food access, rates of GC increased when access to food sources was restricted. Cortisol levels were highest when access to food was constrained. However, because the behavioral and hormonal responses occurred when viewing the stimulus at a distance and preceded the physical presence of the stimulus, we conclude that the anticipation of a competitive situation was sufficient to induce social stress. Contrary to our prediction, targets of aggression did not have higher rates of GC nor did they solicit GC more often than others. Furthermore, higher GC rates did not correlate with a more pronounced decrease in cortisol levels. Not all results obtained in this study supported the predictions concerning the regulatory function of GC on social tension and further research is needed to explore this question. However, the results indicate that the anticipation of competition may be sufficient to induce a costly physiological response, and that high levels of resource competition may have lasting effects on physical stress and stress management. Am. J. Primatol. 71:223–232, 2009. © 2008 Wiley‐Liss, Inc.     

Third-party postconflict affiliation of aggressors in chimpanzees

Postconflict management strategies have been defined as any postconflict interaction that mitigates the negative consequences of the preceding agonistic conflict. Although most studies have investigated postconflict interactions between former opponents or between victims and uninvolved bystanders, interactions between aggressors and bystanders have received much less attention. In this study, we examined a database of 1,102 agonistic interactions and their corresponding postconflict periods in two outdoor-housed groups of captive chimpanzees in order to test the occurrence of postconflict third-party affiliation of aggressors. Our results confirmed the occurrence of appeasement, i.e. postconflict affiliation by a bystander toward an aggressor, but failed to detect the occurrence of postconflict affiliation directed from aggressors toward bystanders. Appeasement rates did not differ according to the sex of the involved individuals. In addition, appeasement occurred more often in the absence of reconciliation than after its occurrence suggesting that appeasement may act as an alternative to reconciliation when the latter fails to occur. Both study groups showed behavioral specificity for appeasement, i.e. context-specific use of certain behaviors, supporting the view that chimpanzees exhibit highly visible explicit postconflict affiliation.     

Asymmetries in postural control and locomotion in chimpanzees (Pan troglodytes)

Posture and locomotion are two of the most primitive and basic motor manifestations of an organism's behavior. Although the restrictions they impose on other motor functions are evident, few studies have considered the possibility of asymmetries in these behaviors in human and nonhuman primates, and how they might impact other asymmetries at higher functional levels. The aim of the current study was to explore in a group of 10 chimpanzees at the Madrid Zoo-Aquarium the degree of asymmetry in four behaviors related to locomotion (walking, ascending, descending, and brachiating) and four behaviors associated with posture (sitting, lying, hanging, and changing postures). Few subjects showed individual preferences, but significant trends in the group for some of the behaviors were found, including right-hand use when initiating quadruped walking, and left-hand use when descending and hanging. Some significant correlations also emerged: a negative one between walking and descending, and a positive one between walking and brachiating and between sitting and changing postures. No correlations were found between locomotor and postural modes. Although we cannot make generalizations on the population level at this time, these findings highlight the importance of considering postural and locomotion factors when studying motor asymmetries in primates.