Imaginary Play in chimpanzees (Pan troglodytes)

Imaginary Play was studied in a group of five signing chimpanzees and it was found that chimpanzees engage in imaginary play similar to that found in human children. Fifteen hours of remote videotapes were analyzed for instances of imaginary play. Behaviors were defined as imaginary play by meeting a predetermined criteria which allowed them to be classified into one of six different categories of imaginary play. Six instances of imaginary play were found and these were classified into the two categories of Animation and Substitution. Observations of imaginary play in other research with chimpanzees were discussed.     

Innovation in wild chimpanzees (Pan troglodytes)

Innovations of behavior have major implications for the concept of culture in animals. Innovation has rarefy been documented in wild animal populations. The chimpanzees of the Taï National Park, Côte d’Ivoire, spontaneously included new patterns of behavior in their repertoire during our study. Some innovations were incorporated into a minority of the group members’ repertoire, whereas others, such as building day nests in trees and on the ground more frequently, became general behaviors. Similarly, new contexts for use of leaf- clipping behavior and novel ways of feeding on some leaves appeared and rapidly became part of the behavior of most group members. The environmental parameters were too stable to explain these new forms of behavior, as most of them took place within 1 month during the same dry season. In a similar way, leaf- grooming acquired a new function in Gombe chimpanzees. A process similar to social conventions could explain the emergence of these new functions for an existing behavior in a way rather similar to human “fashion. ” Three major characteristics of human culture — the absence of individual variations in the performance of the behavior, the general use of the behavior by group members, and the ability to modify the function of a behavior — characterize leaf-clipping in Taï chimpanzees and leaf- grooming in Gombe chimpanzees.     

Molar development in common chimpanzees (Pan troglodytes)

Numerous studies have reported on enamel and dentine development in hominoid molars, although little is known about intraspecific incremental feature variation. Furthermore, a recent histological study suggested that there is little or no time between age at chimpanzee crown completion and age at molar eruption, which is unlikely given that root growth is necessary for tooth eruption. The study presented here redefines growth standards for chimpanzee molar teeth and examines variation in incremental features. The periodicity of Retzius lines in a relatively large sample was found to be 6 or 7 days. The number of Retzius lines and cuspal enamel thickness both vary within a cusp type, among cusps, and among molars, resulting in marked variation in formation time. Daily secretion rate is consistent within analogous cuspal zones (inner, middle, and outer enamel) within and among cusp types and among molar types. Significantly increasing trends are found from inner to outer cuspal enamel (3 to 5 microns/day). Cuspal initiation and completion sequences also vary, although sequences for mandibular molar cusps are more consistent. Cusp-specific formation time ranges from approximately 2 to 3 years, increasing from M1 to M2, and often decreasing from M2 to M3. These times are intermediate between radiographic studies and a previous histological study, although both formation time within cusps and overlap between molars vary considerably. Cusp-specific (coronal) extension rates range from approximately 4 to 9 microns/day, and root extension rates in the first 5 mm of roots range from 3 to 9 microns/day. These rates are greater in M1 than in M2 or M3, and they are greater in mandibular molars than in respective maxillary molars. This significant enlargement of comparative data on nonhuman primate incremental development demonstrates that developmental variation among cusp and molar types should be considered during interpretations and comparisons of small samples of fossil hominins and hominoids.     

Electrocardiogram abnormalities in captive chimpanzees (Pan troglodytes)

Although cardiovascular disease is the leading cause of death in the captive chimpanzee population, little is known about the prevalence and etiology of heart disease in this species. We reviewed the physical exam records of 265 common chimpanzees (Pan troglodytes) for electrocardiogram abnormalities. During the 24-mo period reviewed (August 2003 through August 2005), 34 animals were diagnosed with cardiac arrhythmias consisting of ventricular arrhythmias, supraventricular arrhythmias, conduction disturbances, mixed arrhythmias, and bradycardia. The incidence of cardiac arrhythmia was significantly higher in male animals, chimpanzees 20 to 39 y old, and those with structural heart disease. Incidence of cardiac arrhythmia was not significantly higher in animals with hypertension, hyperlipidemia, or chronic viral infections. During the retrospective period, 7 animals with cardiac arrhythmias died or were euthanized. Mortality was significantly higher in animals with ventricular arrhythmias compared with those without ventricular arrhythmias. We conclude that in the common chimpanzee, age, male gender, and structural heart disease are risk factors for developing cardiac arrhythmias and that ventricular arrhythmias are risk factors for mortality.     

The perception of self-agency in chimpanzees (Pan troglodytes)

The ability to distinguish actions and effects caused by oneself from events occurring in the external environment is a fundamental aspect of human cognition. Underlying such distinctions, self-monitoring processes are often assumed, in which predicted events accompanied by one's own volitional action are compared with actual events observed in the external environment. Although many studies have examined the absence or presence of a certain type of self-recognition (i.e. mirror self-recognition) in non-human animals, the underlying cognitive mechanisms remain unclear. Here, we provide, to our knowledge, the first behavioural evidence that chimpanzees can perform self/other distinction for external events on the basis of self-monitoring processes. Three chimpanzees were presented with two cursors on a computer display. One cursor was manipulated by a chimpanzee using a trackball, while the other displayed motion that had been produced previously by the same chimpanzee. Chimpanzees successfully identified which cursor they were able to control. A follow-up experiment revealed that their performance could not be explained by simple associative responses. A further experiment with one chimpanzee showed that the monitoring process occurred in both temporal and spatial dimensions. These findings indicate that chimpanzees and humans share the fundamental cognitive processes underlying the sense of being an independent agent.     

Not all chimpanzees (Pan troglodytes) show self-recognition

When a chimpanzee is presented with a mirror it initially responds with social behavior directed toward the reflection. After several hours of exposure to the mirror the social behavior decreases and the mirror is used to guide self-directed responses to previously unobservable parts of the body such as the face. When a distinctively colored mark is unobtrousively applied to the chimpanzee's face and the chimpanzee touches the mark while observing itself in the mirror, this behavior is said to indicate self-recognition. Such self-recognition has been considered to be a robust phenomenon in chimpanzees, with self-directed and mark-directed behaviors both appearing in all socially-housed adult chimpanzees tested. In our study 11 chimpanzees were given mirror exposure and tested with the mark test. Only one of the 11 chimpanzees touched the mark during test, although several showed self-directed behavior using the mirror to guide their movements. Such experimental factors as mirror size, position, or temporal spacing of the mirror exposure, and such subject variables as age, sex, previous social experience, and subspecies were insufficient to explain the difference between the present and previous findings. We suggest that there are individual differences in mirror recognition behavior in chimpanzees, and that further consideration of the factors contributing to this phenomenon, including the development of additional tests for self-recognition, is needed.     

Normal vaginal flora in chimpanzees (Pan troglodytes): qualitative and quantitative study

Lactobacilli are the predominant microorganisms in the vaginal flora of human beings, and are known to play an important role in protecting them from genital infections. On the other hand, the composition of the vaginal flora differs among laboratory animal species, and lactobacilli are not the predominant vaginal microorganism in many laboratory animals. We speculated that the vaginal flora of chimpanzees would be more similar to those of human beings than to those of other animal species, because chimpanzees are phylogenetically close to human beings, and their reproductive physiology is similar to that of human beings. To clarify our speculation, we examined the development of the vaginal flora in chimpanzees (Pan troglodytes). Streptococci, lactobacilli, and members of the family Bacteroidaceae were the most predominant bacteria in the vagina of mature chimpanzees (9 to 22 years old). During development of the vaginal flora of chimpanzees, the total number of bacteria increased with age and reached a plateau just before sexual maturity (5 to 7 years of age; juvenile period). Lactobacilli were already one of the predominant bacteria before sexual maturity. In mature chimpanzees, the total number of bacteria (aerobes and anaerobes) in the vagina was highest during the swelling phase of the menstrual cycle. During the swelling phase in mature chimpanzees, streptococci, lactobacilli, and Bacteroidaceae were the most frequently isolated (100%) organisms, and the total number of organisms recovered from vaginal specimens from these three groups was the highest. In mature chimpanzees in which the number of bacteria was the highest, lactobacilli were the predominant bacteria. Taken together, these results suggest that these three bacterial groups (streptococci, lactobacilli, and Bacteroidaceae) are indigenous to the vagina of chimpanzees, and chimpanzees would be the most suitable laboratory animals for studying the role of lactobacilli in the vagina of human beings.     

Natural genital herpesvirus hominis infection in chimpanzees (Pan troglodytes and Pan paniscus)

Type 2 Herpesvirus hominis was isolated from pustulovesicular lesions on the external genitalia of two chimpanzees. Histopathologic examination of biopsy specimens from both animals revealed typical herpetic changes which included necrosis, superficial ulceration acute inflammatory cell infiltration, multinucleated syncytial giant cells, and intranuclear inclusions. Large numbers of herpes-type viruses were demonstrated by electron microscopy in biopsy specimens from both animals. Serologic studies also demonstrated infection of these animals with Herpesvirus hominis.     

Design complexity in termite-fishing tools of chimpanzees (Pan troglodytes)

Adopting the approach taken with New Caledonian crows (Corvus moneduloides), we present evidence of design complexity in one of the termite-fishing tools of chimpanzees (Pan troglodytes) in the Goualougo Triangle, Republic of Congo. Prior to termite fishing, chimpanzees applied a set of deliberate, distinguishable actions to modify herb stems to fashion a brush-tipped probe, which is different from the form of fishing tools used by chimpanzees in East and West Africa. This means that ‘brush-tipped fishing probes’, unlike ‘brush sticks’, are not a by-product of use but a deliberate design feature absent in other chimpanzee populations. The specialized modifications to prepare the tool for termite fishing, measures taken to repair non-functional brushes and appropriate orientation of the modified end suggest that these wild chimpanzees are attentive to tool modifications. We also conducted experimental trials that showed that a brush-tipped probe is more effective in gathering insects than an unmodified fishing probe. Based on these findings, we suggest that chimpanzees in the Congo Basin have developed an improved fishing probe design.     

Videotapes as enrichment for captive chimpanzees (Pan troglodytes)

The effectiveness of showing videotapes to captive chimpanzees as an environmental enrichment was quantitatively tested. The responses of 10 subjects (3 adult males and 7 adult females) to videotapes of chimpanzees engaging in a variety of behaviors, to videotapes of other animals and humans, and to television programs were compared. Data collection consisted of 20-minute, continuous sampling tests while various videotapes were shown. A total of 400 tests were conducted. Multivariate analysis of variance was applied to measure differences in the duration of eight categories of behavior across videotapes of varying content. No general behavioral differences in response to the tapes based on sex or housing were revealed. However, with the behavior of monitor-watching analyzed alone, we found that individually housed subjects watched the videotapes more than socially housed subjects. When viewing time was averaged across all videotapes, the chimpanzees watched the monitor a mean of 38.4% of the time available. The chimpanzees' behavior varied significantly only when they were watching the videotapes of various human and chimpanzee activities and not when watching a blank screen. A Pearson's correlation indicated that subjects habituated to repeated presentations of the videotapes, although the effect was small numerically. Although this type of enrichment did not extensively alter behavior, it did occupy a significant portion of the subjects' activity budget; thus, the amount of time spent watching the video stimuli indicated that videotapes may be a useful enrichment for captive chimpanzees. Zoo Biol 19:541-551, 2000. Copyright 2000 Wiley-Liss, Inc.     

Evidence for social referencing in young chimpanzees (Pan troglodytes)

A captive chimpanzee group was observed in order to determine the extent to which the social interactions of the infants and juveniles (18-50 months) were affected by their mothers' relationships with other adult group members. It was found that the young chimpanzees initiated more interactions with adults who interacted more with their mothers. A vast majority of those interactions occurred at significant distances from the mother. It is argued that these data imply a social-cognitive ability in young chimpanzees closely related to the human infant's ability to use its mother in 'social referencing'.     

What chimpanzees (Pan troglodytes) learn in a cooperative task

To examine the development of cooperation in a captive group of chimpanzees (Pan troglodytes), we designed an apparatus which required the simultaneous traction by two animals to get a reward. Two chimpanzees, an adult male and an infant female in a group of six, produced most of the successful responses (pulling on two handles simultaneously). Visual behavior was used to try to determine what chimpanzees learned about the cooperative task. Propositions were made to investigate what kind of learning could be attributed to chimpanzees and were confronted with results. Both subjects learned the link between the presence of fruits on the apparatus and the possibility of getting a fruit. They also learned the importance of the partner at the apparatus to make a successful response. Only the adult male learned to take into account the behavior of the partner at the apparatus before pulling a handle. From a methodological point of view, the glances made by the animals can constitute a useful behavioral indicator of what the subjects learned in a given social situation.     

Behavioral responsiveness to strangers in young chimpanzees (Pan troglodytes)

Earlier studies of responses of primate infants to strangers were frequently confounded with separation from the attachment figure and/or removal to an unfamiliar setting. In the present study, young chimpanzees were tested in a familiar setting with a human female caretaker who served as an attachment figure. Two strangers were introduced: a human female, approximately the size of the caretaker, who refrained from initiating interactions and a larger human male, who repeatedly approached and initiated contact with the subjects. A somewhat more intense response was elicited by the larger, more assertive male stranger, but neither stranger elicited severe distress. The behavior of the chimpanzees is better described as wary, rather than fearful. These results may be attributable to the continued presence of the caretaker in contrast to previous studies reporting severe distress to strangers when chimpanzees are tested alone. A wariness of strangers in nursery-reared chimpanzees may develop by 6 months of age and may diminish somewhat by 2 years of age under the conditions of this study.     

Controllability in environmental enrichment for captive chimpanzees (Pan troglodytes)

This study considers the use of nonsocial environmental enrichment by captive chimpanzees at the Primate Foundation of Arizona. The goal was to determine whether a relationship existed between controllability of enrichment items by captive chimpanzees and frequency of use. The study measured controllability, the ability of nonhuman animals to alter aspects of their environment by the potential destructibility of the enrichment item. This study examined additional factors that may affect enrichment use: individual age, sex, rearing history, social group composition, and availability of outdoor access. The chimpanzees in the study used destructible items—the enrichment category with the highest level of controllability—more than indestructible items across all age, sex, and rearing classes. Thus, controllability seems to be an important factor in chimpanzee enrichment. Younger individuals and groups with outdoor access used enrichment more than did older individuals and groups with indoor-only access. Individual sex, rearing history, and social group composition had minimal effects on enrichment use. These results support the importance of control to captive chimpanzees and further enable captive management to customize enrichment programs to the needs of particular animals.     

Social behavior and genital swelling in pregnant chimpanzees (Pan troglodytes)

Previous reports of increased sexual activity associated with genital swelling during chimpanzee pregnancy have suggested that the tumescent pregnant female may be regarded as a regularly cycling estrous female. Analysis of additional data from seven pregnant and eight cycling females, however, indicated that tumescent pregnant females differed from cycling females in some social interactions with males. As in earlier reports, pregnant females were more receptive to male-initiated copulatory bouts. In addition, pregnant females initiated grooming bouts with males much less frequently than did cycling females, and males spent less time grooming and less frequently inspected the genitals of pregnant than cycling cagemates. This tendency to decrease social interaction during pregnancy is consistent with field reports that pregnant chimpanzees prefer the company of their offspring and other noncycling females. In addition, the genital swelling patterns of 40 chimpanzee pregnancies were analyzed. Length of gestation averaged 231.5 days. Although a high degree of individual variation existed, females were swollen an average of 41% of the days observed, yet reached maximum swelling only 8.7% of the time. Genital swelling was less frequent during the third trimester, averaging 50.5, 47.9, and 22.6% tumescent days in each trimester. Young pregnant females showed maximum swelling more often than did older females. This finding is discussed with respect to field reports of intercommunity transfer of young, tumescent female chimpanzees.     

Comparing infant and juvenile behavior in bonobos (Pan paniscus) and chimpanzees (Pan troglodytes): a preliminary study

The dichotomy between the two Pan species, the bonobo (Pan paniscus) and chimpanzee (Pan troglodytes) has been strongly emphasized until very recently. Given that most studies were primarily based on adult individuals, we shifted the “continuity versus discontinuity” discussion to the infant and juvenile stage. Our aim was to test quantitatively, some conflicting statements made in literature considering species differences between immature bonobos and chimpanzees. On one hand it is suggested that infant bonobos show retardation in motor and social development when compared with chimpanzees. Additionally it is expected that the weaning process is more traumatic to chimpanzee than bonobo infants. But on the other hand the development of behaviors is expected to be very similar in both species. We observed eight mother–infant pairs of each species in several European zoos. Our preliminary research partially confirms that immature chimpanzees seem spatially more independent, spending more time at a larger distance from their mother than immature bonobos. However, the other data do not seem to support the hypothesis that bonobo infants show retardation of motor or social development. The development of solitary play, environmental exploration, social play, non-copulatory mounts and aggressive interactions do not differ between the species. Bonobo infants in general even groom other group members more than chimpanzee infants. We also found that older bonobo infants have more nipple contact than same aged chimpanzees and that the weaning process seems to end later for bonobos than for immature chimpanzee. Additionally, although immature bonobos show in general more signs of distress, our data suggest that the weaning period itself is more traumatic for chimpanzees.     

Features of meat digestion by captive chimpanzees (Pan troglodytes)

The pronounced carnivory of many human populations contrasts sharply with feeding habits of other Hominoidea. Of extant great apes, only chimpanzees (Pan spp.) actively seek out vertebrate prey, but meat is only a minor portion of their diet. Some accounts suggest that wild chimpanzees digest prey inefficiently. To investigate the capacity of chimpanzees to digest meat, feeding trials were carried out on three captive chimpanzees (Pan troglodytes) using a fixed amount of nonpurified diet with and without a predetermined amount of boned cooked chicken. The results showed no significant differences in the rate of passage of digesta and digestion of diets with and without chicken. Meat ingestion did not change the nitrogen (N) concentration of feces or the total amount of N defecated. Visual inspection of fecal matter showed no evidence of undigested meat. Taken together, the results indicate that chimpanzees are able to digest meat of the type and quantity consumed during these trials.     

Synchrony of estrous swelling in captive group-living chimpanzees (Pan troglodytes)

Recent findings suggest that menstrual synchrony may occur in humans and is more dependent on factors of close personal association than simple prox imity. Part I of this study analyzed the estrous cycles of 10 captive group living chimpanzees (Pan troglodytes) housed in two separate groups in the same building. Each group contained at least one male. The mean discrepan cies for (1) the approximate day of ovulation (last day of maximum swelling) and (2) the day of onset of estrous swelling were determined for all possible pairs within groups and compared to the mean discrepancies of pairs between groups. With respect to the probable day of ovulation, the mean discrepancy of animals caged together was 8.0 days and that of animals caged separately was 10. 7 days. Comparing the onset of estrous swelling, the mean between groups was 10.4 days, whereas females caged together averaged only 5.7 days. A repeated-measures analysis of variance indicated a significant group effect in the onset of estrous swelling (F = 5.68,p < 0.05), whereas there was no significant difference between the animals caged together and those caged separately with respect to the last day of maximum swelling (F = 1.37, NS). Part II of the study compared the onset of the first postpar tum estrus (PPE) to the onset of estrus in cagemates. During a 6-year period, 13 females exhibited 33 PPEs while in social contact with cycling females. Of 86 paired comparisons, the PPE onset of 43 (50%) occurred with in 6 days of the swelling onset of a cagemate. The results suggest that, as in other species (including humans), the social contact and time spent together influence the synchrony of chimpanzee estrous cycles, especially in terms of swelling onset. The synchrony of estrous swelling and possible mediation by olfactory and/or hormonal cues are discussed as they relate to chimpanzee social structure and female reproductive strategy.